(World Crop Series) W. J. Florkowski (Auth.), J. Smartt (Eds.) - The Groundnut Crop - A Scientific Basis For Improvement-Springer Netherlands (1994) PDF

THE GROUNDNUT CROP
W orld Crop Series
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Its scientific basis
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THE
GROUNDNUT
CROP
A scientific basis for improvement
Edited by
J. Smartt
Department of Biology
Southampton University, UK
IDIlI SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.

© Springer Science+Business Media Dordrecht 1994
Softcover reprint ofthe hardcover lst edition 1994
Originally published by Chapman & Hali
ISBN 978-94-010-4315-1 ISBN 978-94-011-0733-4 (eBook)

DOI 10.1007/978-94-011-0733-4
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Contents
List of contributors x
Series foreword xv
Preface xvii
1 Groundnut production and trade 1

Wi. Florkowski
1.1 Production, area harvested and yield 1
1."-'r) Stocks and prices 15
1.3 International trade 17
1.4 Concluding comments 21
Acknowledgements 22
References 22
2 The origin and history of the groundnut 24

R. O. Hammons
2.1 Introduction 24
2.2 Prehistoric groundnut agriculture 25
2 ..)r, Post-Columbian historical narratives 26
2.4 Indian vernacular names 31
2.5 Origin of groundnut 31
2.6 Dispersal 33
2.7 Ethnography 35
2.8 Early industrial developments 36
References 39
3 Botany - morphology and anatomy 43

V. Ramanatha Rao and UR. Murty
3.1 Introduction 43
3.2 Description of the genus 43
3.3 Description of the species 44
3.4 Infraspecific differentiation and morphology of
Arachis hypogaea 53
3.5 Morphology and development 55
VI Contents
3.6 Anatomy 75
3.7 Concluding remarks 88
References 89
4 Biosystemics and genetic resources 96
A.K. Singh and c.E. Simpson
4.1 Introduction 96
4.2 Nomenclature 97
4.3 Systematics 99
4.4 Phylogeny 107
4.5 Genetic resources 109
4.6 Gene pools 110
4.7 Centres of origin 112
4.8 Centres of diversity 113
4.9 Collections 118
4.10 Genetic variability 122
4.11 Conservation of germplasm 129
4.12 Utilization 131
4.13 Conclusions and perspectives 132
References 133
5 Reproductive biology and development 138
P. Coolbear
5.1 Flower development, pollination and fertilization 138
5.2 Fruit development 140
5.3 Seed development 144
5.4 The biochemistry of seed development 149
5.5 Seed dormancy 153
5.6 Losses of potential yield 155
5.7 Production conditions and seed quality 158
5.8 Quality factors and the process of seed deterioration 162
5.9 Some final conclusions 167
References 167
6 The composition and nutritive value of groundnut kernels 173
G. P. Savage and 1.1. Keenan
6.1 Introduction 173
6.2 Proximate composition 173
6.3 Oil and fatty acids 174
6.4 Protein 183
6.5 Carbohydrate 196
6.6 Vitamins 200
6.7 Mineral composition 201
6.8 Other components 202
6.9 Conclusions 204
Acknowledgements 205
References 205
Contents Vll
7 Mineral nutrition 214

G.J. Gascho and 1. G. Davis
7.2 Calcium (Ca) 215
7.3 Nitrogen (N) 226
7.4 Phosphorus (P) 228
7.5 Potassium (K) 230
7.6 Magnesium (Mg) 231
7.7 Sulphur (S) 233
7.8 Micronutrients 234
7.9 Current emphasis and research needs 244
References 245
8 Nitrogen fixation 255
1. Sprent
8.2 Arachis as a nodulating genus 257
8.3 The rhizobial component 260
8.4 Functional nodules 263
8.5 Measurement of nitrogen fixation in the field 268
8.6 Effects of environmental stress in nodulation and
nitrogen fixation 272
8.7 Prospects for improvement 275
References 276
9 Groundnut water relations 281
G. C. Wright and R. C. Nageswara Rao
9.2 Groundnut water status 281
9.3 Effects of water deficit on some physiological and
morphological processes 284
9.4 Effect of water deficits on root and shoot growth 287
9.5 Factors affecting evapotranspiration 302
9.6 Yield responses to evapotranspiration and
transpiration 305
9.7 Ameliorating the effects of water deficits on yield 309
9.8 Conclusions 324
References 325
10 Diseases 336
K.J. Middleton, S. Pan de, S.B. Sharma and D.H. Smith
10.1 Diseases caused by bacteria 336
10.2 Diseases caused by viruses and mycoplasmas 339
10.3 Diseases of foliage, caused by fungi 346
Vlll Contents
10.4 Diseases of stems, roots and pods, caused by fungi 359
10.5 Diseases caused by nematodes 372
References 378
11 Groundnut pests 395

l.A. Wightman and G. V. Ranga Rao
11.2 The insects 399
11.3 Applied ecology and economics 415
11.4 Host plant resistance 430
11.5 Natural enemies of groundnut insects 441
11.6 Insecticides 450
11.7 Cultural control 454
11.8 Research related to insect pests of stored groundnut 461
11.9 Integrated control of ground nut pests 463
11.10 Conclusions 467
References 469
12 Industrial utilization and processing 480

l.J.K.B. Asiedu
12.1 Utilization 480
12.2 Processing 482
12.3 Harvesting, stripping and drying 483
12.4 Storage 485
12.5 Shelling or decorticating 486
12.6 The processing of groundnuts into peanut butter 487
12.7 The processing of roasted and salted groundnuts 491
12.8 The processing of peanut candies and confections 494
12.9 Oil expression 497
References 508
13 Mycotoxins in groundnuts, with special reference to aflatoxin 509

l.l. Keenan and G.P. Savage
13.2 Natural occurrence of aflatoxins 510
13.3 Preharvest contamination 511
13.4 Post-harvest contamination 513
13.5 Presence in foodstuffs 514
13.6 The chemistry and metabolism of aflatoxin 517
13.7 Aflatoxicosis in animals and poultry 522
13.8 Aflatoxin and human disease 524
13.9 Analytical methods for the detection of aflatoxins 528
Contents IX
13.10 Overcoming aflatoxin contamination 535

References 541
14 Groundnut breeding 552
T.G. Isleib, J.C Wynne and S.N. Nigam
14.2 Variability in groundnut germplasm 553
14.3 Breeding methods 574
14.4 Regional progress 575
14.5 Accomplishments and future efforts 615
References 617
15 Utilization of Arachis species as forage 624
B. G. Cook and I. C Crosthwaite
15.2 Arachis hypogaea 625
15.3 Wild species 633
15.4 Breeding 655
15.5 Conclusion 656
References 657
16 The groundnut in farming systems and the rural economy
- a global view 664
1. Smartt
16.1 Farming systems 664
16.2 Cultivation systems 666
16.3 Adaptive research 675
References 699
17 The future of the groundnut crop 700
J. Smartt
17.1 Product diversity 700
17.2 Farming groundnut 702
17.3 Improving the groundnut 709
17.4 The role of biotechnology 714
17.5 Agricultural constraints, research and the future 716
17.6 A new philosophy 719
References 719
Index 721
Contributors
J.J.K.B. Asiedu
The African Regional Centre for Technology
BP 2435
A venue Djily Mbaye
Dakar
Senegal
B.G. Cook
Senior Pasture Agronomist
Queensland Department of Primary Industries
Cnr Cartwright Road and Louisa Street
PO Box 395
GYMPIE
Queensland 4570
Australia
P. Coolbear
Seed Technology Centre
Massey University
Palmerston North
New Zealand
I.C. Crosthwaite
Extension Agronomist
Kingaroy
Queensland
Australia
J.G. Davis
Department of Crop and Soil Sciences
University of Georgia
Coastal Plain Experiment Station
Tifton
GA 31793
USA
Contributors xi
W.J. Florkowski
Associate Professor
The University of Georgia
College of Agricultural and Environmental Sciences
Department of Agricultural and Applied Economics
Georgia Experiment Station
Griffin
GA 30223-1797
USA
G.J. Gascho
Department of Crop and Soil Sciences
University of Georgia
Coastal Plain Experiment Station
Tifton
GA 31793
USA
R.O. Hammons
Hammons Consultancy
1203 Lake Drive
Tifton
Georgia 31794-3834
USA
T.G. Isleib
Department of Crop Science
North Carolina State University
Raleigh
NC 27695-7620
USA
J.1. Keenan
Department of Surgery
Christchurch School of Medicine
University of Otago
New Zealand
K.J. Middleton (deceased)

1. Bjelke Petersen Research Station
POBox 23
Kingaroy 4610
Queensland
Australia
xu Contributors
U.R. Murty
Director
National Research Centre for Sorghum
Rajendranagar
Hyderabad
Andhra Pradesh 500 030
India
R.C. Nageswara Rao
Legumes Program
International Crops Research Institute for the Semi-Arid
Tropics (ICRISAT)
Patancheru
India
S.N. Nigam
Principal Groundnut Breeder
Tropics (ICRISAT)
Patancheru
India
S. Pan de
Saudi Agricultural Development Co. (INMA)
P.O. Box 148
Alkamasen
Wadi-Al-Dawasir Il9I
Kingdom of Saudi Arabia
V. Ramanatha Rao
International Board for Plant Genetic Resources
Regional Office for Asia, the Pacific
and Oceania
7th storey, RELC Building
30 Orange Grove Road
Singapore 1025
G.V. Ranga Rao
Legumes Program
Tropics (ICRISAT)
Patancheru
India
Contributors xiii
G.P. Savage
Department of Biochemistry and Microbiology
Lincoln University
Canterbury
New Zealand
S.B. Sharma
Tropics (ICRISAT)
Patancheru, Andhra Pradesh 502 324
fndia
C.E. Simpson
Texas Agricultural Experiment Station
Texas A & M University
Stephenville
Texas 76401
USA
A.K. Singh
Genetic Resources Unit
International Crops Research for the Semi-Arid
Tropics (fCRISA T)
Patancheru
India
J. Smartt
School of Biological Sciences
Department of Biology
Biomedical Sciences Building
University of Southampton
Bassett Crescent East
Southampton S09 3TU
UK
D.H. Smith
Tropics (fCRISAT)
Patancheru
India
J. Sprent
Department of Biological Sciences
The University
Dundee DDI 4HN
UK
xiv Contributors
J.A. Wightman
Legumes Program International Crops Research Institute for the
Semi-Arid Tropics (ICRISAT)
Patancheru
India
G.e. Wright
1. Bjelke Petersen Research Station
Kingaroy
Qld 4610
Australia
J.e. Wynne
Head
Department of Crop Science
North Carolina State University
Raleigh
NC 27695-7620
USA
Series foreword
This book is a most welcome addition to the World Crop Series. For
although in terms of world figures groundnut annual production (in shell)
at 23 506 000 tonnes is second to and well below that of soyabean at
114011 000 tonnes of grain, it is of special importance in developing
countries. Thus about ten times as much of the groundnut crop is produced
in developing countries than in developed countries, whereas well over half
the production of soyabean is in developed countries (FAD Yearbook,
1992). Moreover, the groundnut crop has a special role in sustaining the
agricultural systems of the semi-arid tropics where, because of the stressful
climates, farming is particularly difficult.
This series recognizes that, because the disciplines contributing to crop
improvement are becoming more specialized, it is increasingly difficult for
the specialists involved to understand each other's language and percep-
tions. It is not only a problem for those active in research but is perhaps an
even greater difficulty for those who are involved in agricultural develop-
ment, extension and education to obtain a critical, balanced and up-to-date
view of the scientific advances that are contributing to crop improvement.
At the same time it is evident from the way in which research priorities
and programmes are being addressed internationally that increasing
emphasis is being placed on multidisciplinary projects. Hence, in spite of
increased specialization, the need for scientists to understand each other
and integrate their efforts is also increasing. The multidisciplinary
approach is evident, for example, in the project-based system of organiz-
ation and management that has been recently introduced in the
International Crops Research Institute for the Semi-Arid Tropics (ICRI-
SAT). This is the institute within the system of eighteen International
Centres of the Consultative Group for International Agricultural Research
which has been given a special responsibility to work on the groundnut
crop and also to conserve the world germplasm collection of this species
and its wild relatives. It is not surprising, therefore, that several of the
authors of this book are ICRISAT scientists. But interest in the crop is not
restricted to the semi-arid tropics; and while the major production regions
are India, China and Africa (in that order) it is also important elsewhere,
XVI Series foreword
especially in the USA. Scientific expertise in the crop extends even wider
and the authors who were invited to contribute to this book by the editor
reflect the considerable international interest in groundnuts.
I believe the result of this combined effort will become an essential
source of ideas and information for anyone who takes more than a passing
interest in this crop. While making no scientific compromises, the language
is meant to be comprehensible to anyone with some understanding of
biological and agricultural sciences. It should therefore be a considerable
aid to the multidisciplinary approaches that are now a feature of agricul-
tural research and development.
E.H. Roberts
University of Reading
Preface
The event which threw the 'Groundnut Crop' into sharpest relief was
undoubtedly the monumental fiasco of the East African Groundnut
Scheme, conceived in ] 946. What had been an interesting botanical curio-
sity and the source of useful foods, edible oils and feeding stuffs became
the butt of innumerable comedians and to be in any way associated with
the crop in the 1950s was to be a sitting target for wits and humorists of
every stripe. To be informed within 24 hours of arrival in Northern
Rhodesia that I was to devote my attention and energies to this crop made
me feel that I had drawn the shortest of short straws. It was some while
before gloom and despondency lifted but by the end of my first crop season
I realized that in fact I had not been given the metaphorical poisoned
chalice but something that was to become an abiding interest, which has
remained fresh over a period of 40 years.
Three events in that first season (1954-5) launched me on a course which
would over the years give me enormous job satisfaction. Stephen Hoyle of
the Nyasaland Department of Agriculture (a groundnut enthusiast if ever
there was one) made available to me his own variety collection. Early in
1955 I received from Queensland, in response to a request for groundnut
material, several selections of a Bolivian groundnut landrace named Mani
Pintar which were to prove extremely interesting. The final gift of the fairy
godmother was a copy of The Peanut - the Unpredictable Legume, which
became my groundnut bible during my spell in Africa, as it did for many
others in the groundnut (or peanut) world. It was so successful and highly
regarded that its successor Peanuts - Culture and Uses did not appear until
1973 - a lapse of 22 years. In the meantime, however, two useful if modest
publications were produced in 1967: the PANS (Pest Articles and News
Summaries) Manual No.2, Pest control in GROUNDNUTS (published
by the Overseas Development Agency in the UK) and Groundnut
ProductionlGroundnut Research 1954-61 published in Lusaka by the
Government Printer, Zambia.
Advances in production technology and research continued at an acce-
lerating pace and in 1982 the American Peanut Research and Education
Association (now Society) published Peanut Science and Technology, a
XVlll Preface
successor to its earlier volume, Peanuts - Culture and Uses. This organiz-
ation has since 1974 published its own journal Peanut Science which
provides a useful vehicle for publication on any aspect of investigation
related to the crop. Prior to that date, APREA had published its own
untitled journal for a number of years. In addition, the Proceedings of its
annual meetings are invaluable sources of information - going back to the
early 1960s if one also includes those of its predecessor, the Peanut
Improvement Working Group.
The first major work on the groundnut to emerge from the developing
world was published in 1988 by the Indian Council of Agricultural
Research with a similar format and organization of material to the APREA
publications. This very valuable and substantial piece of work is concerned
primarily with the Indian context while the corresponding APREA publi-
cations address the North American situation. Thus there appears to be
scope for a broader approach to the crop, incorporating experience with
the groundnut crop in the rest of the world. This was the motivating force
behind the present volume, which attempts where possible to break new
ground with the presentation of authoritative reviews such as that of
Professor Janet Sprent as well as broad geographic coverage of other topics
such as breeding and the control of pests and disease.
Any editor setting about the task of assembling a cast of authors does so
with a set of ideals in mind, well knowing whom to recruit. The process is
beset with trials and disappointments and this volume has had its fair share
or more of these. It is with the deepest regret that the death of a major
contributor, Keith Middleton, must be recorded. The chapter on Diseases
had been fully drafted by the time of his death and was completed by his
colleagues at ICRISAT. I count myself privileged and honoured to have met
him in the year before his untimely death. He will be sorely missed.
The current depressed state of the world economy has not been without its
effects on contributors who had agreed in good faith to produce chapters but
in the event were unable to do so. An individual threatened by the horrors of
'rationalization' may not be in the best condition to discharge obligations
gladly accepted in happier times. A somewhat fuller coverage had been
anticipated but in fairness to contributors who had met the deadlines the
decision was taken to proceed, with the editor making an attempt to fill as far
as possible the lacunae left by the contributions which failed to materialize.
Acknowledgement must be given to ICRISAT's work on the groundnut
crop, guided by Dr J. G . Ryan, its present director, and his predecessor Dr
L.D. Swindale. The encouragement given by Dr Y.L. Nene as Deputy
Director General is acknowledged gratefully. The experience of editing
this work has been very rewarding and has given me a pretext for making
contact with those whose work I have long admired but whom otherwise I
might never have approached.
J. Smartt
Southampton, October, 1993
CHAPTER 1
Groundnut production and trade

W.]. Florkowski
This chapter focuses on major issues and implications of groundnut pro-

duction, processing, distribution and consumption, discussion of yields and
area harvested, world prices and trade illustrates strong and weak links in
the world groundnut sector.
Revenue generated from groundnut sales amounts to billions of dollars
annually and sales for domestic use are important to regional and national
economies. Groundnut is a high value product and an important source of
cash to farmers in less developed countries, allowing the purchase of other
foods and industrial goods.
Groundnut oil is a major vegetable oil in many parts of the world and its
sale to domestic and foreign consumers generates revenue. Its production
by mills crushing locally grown groundnut provides jobs and income for
numerous workers. By-products, such as groundnut cake and meal,
directly or indirectly improve nutrition.
Export sales of groundnut, groundnut oil and cake can increase the
income of groundnut farmers and processors. Groundnut exporters include
a number of countries in Africa, Asia and the Americas but the three
major exporters in the 1980s were the USA, Argentina and China. An
increase in yield and in the area harvested led to increased production and
exports from China and Argentina. Many countries with increasing yields
concentrate on satisfying a strong domestic demand but are likely to export
in years of world shortage, taking advantage of a high price.
1.1 PRODUCTION, AREA HARVESTED AND YIELD
Several factors determine changes in yield, area harvested and level of

production. Applied technology, better management and the weather
directly influence yield. The area under crop is determined by groundnut
price, expected profits from groundnut relative to other crops, and the
The Groundnut Crop: A scientific basis for improvement. Edited by 1. Smartt. Published in
1994 by Chapman & Hall, London. ISBN 0 412 408201.
2 Groundnut production and trade
inputs available. Both policy and weather influence yield and area har-
vested. Policy is a factor that represents a broad spectrum of macroecono-
mic variables. Foreign exchange needs frequently lead to a government
decision to increase production of an exportable commodity, as was the
case in China. Food needs of growing populations could accelerate adop-
tion of higher yielding cultivars.
In the following sections, discussion of the groundnut industry on each
continent centres on selected producing countries which illustrate changes
in the groundnut sector of specific regions. Trends in area harvested and
production among countries reveal the changing importance of the dom-
estic groundnut sector.
For several countries, continuous information on the industry is not
available for the entire period (1960-1990). Reasons for gaps in the
statistics include creation of sovereign states, social unrest and military
conflicts. For the purposes of this presentation, however, the available
information is adequate.
1.1.1 Groundnut production

Groundnut production is important to several countries with large popu-
lations, in which groundnut plays a role as a food crop. The volume of
production reflects the amount of resources allocated to groundnut and its
importance to a particular economy. Table 1.1 compares the average
output for selected countries over three decades (1960s-1980s), and Table
1.2 highlights changes in production levels on a percentage basis. Some of
the more obvious trends are as follows.
(a) North America

In Mexico, production decreased in the 1970s but recovered in the 1980s
with an increase of 22%. In the USA, a production increase of 62%
between the 1960s and the 1970s reflected technological gains.
(b) South America

In the large groundnut countries, production has been decreasing. The
opposite trend is true in many smaller countries. Brazil was the largest
producer in South America in the mid 1960s and the 1970s but its output
decreased by 53% during the 1980s in comparison with the 1970s. Since
1986, Argentina has been consistently the largest producer in South
America: production increased by 23% between the 1960s and the 1970s
but decreased by 30% during the 1980s. The overall decrease in production
in South America was the result of a smaller area being sown to groundnut
in Brazil and Argentina.
Production, area harvested and yield 3
(c) Africa
In Africa, countries that reported decreased groundnut production out-
numbered those with an increase in the 1980s. The average for individual
countries varied widely over the three decades but the continent's total
increased slightly, having declined during the 1970s.
In West Africa, there was a decrease in major producing countries
during the 1980s over the 1960s, although output was fairly stable for the
region as a whole and it is noticeable that the Ivory Coast nearly doubled
its output in the 1980s, while in Benin production increased by 134%
between the 1960s and the 1980s.
In eastern Africa, overall production increased over the period 1960-
1989. In the Sudan, the increase was by 136% during the 1970s, but
Uganda's average output in the 1980s dropped to 37% below its 1960s
level.
In central Africa, Zaire's output increased by 89% over the three
decades. In southern Africa, the trend was for a decrease in production of
groundnut.
(d) Asia
All major producers in Asia increased their average output over the three
decades. In south-west Asia, India's production climbed steadily over the
period, increasing by 25%. In south-east Asia (excluding Vietnam), pro-
duction increased by 176% between the 1960s and 1980s. In Myanmar, the
average output rose steadily, increasing by 16% during the 1970s and a
further 36% during the 1980s. Indonesia more than doubled its average
output.
East Asia's output grew mainly because of China, which recorded a 90%
increase in average annual groundnut production between the 1970s and
the 1980s. However, output fell in Taiwan and did so more dramatically in
Japan, where it declined by 63% between the 1960s and the 1980s.
(e) Europe
The groundnut is of marginal importance to European farmers. The grow-
ing conditions in most European countries are unsuitable for its
production. In several countries where it is grown, there may be a com-
parative advantage in production of other crops. Relatively high yields
have not compensated for the small area and, as a result, the production of
groundnuts has been low in most of Europe, seldom exceeding 10 000 MT
in anyone country. Turkey has been the largest producer: in the 1960s, it
averaged 25 400 MT annually, increasing this level by 77% in the 1970s and
another 34% in the 1980s.
MT = (onnes.
TABLE 1.1 Average groundnut production in selected countries (1000 tonnes)
Change Change
Country 1960-1969 1970-1979 1970s/1960s 1980-1989 1980sl1970s
North America
Mexico 89.7 59.5 -30.2 72.9 13.4
USA 992.9 1606.8 613.9 1669.7 62.9
South America
Argentina 325.8 402.1 76.3 281.8 -120.3
Brazil 588.1 582.8 -5.3 272.6 -310.2
Colombia NA 2.6 NA 4.4 2.2
Ecuador 10.9 10.3 -0.6 9.8 -0.5
Paraguay 13.6 17.5 3.9 36.0 18.5
Uruguay 4.8 2.6 -2.2 1.2 -1.4
Venezuela 2.4 19.6 17.2 15.4 -4.2
Africa
West Africa 285.5 209.0 -76.5 184.1 -24.9
Benin 24.0 49.9 25.9 56.2 6.3
Burkina Faso 114.1 118.0 3.9 123.7 5.7
Cameroon 120.5 123.5 3.0 185.6 62.1
Ghana 59.6 74.8 15.2 63.7 -9.1
Gambia 100.3 123.1 22.8 114.9 -8.2
Ivory Coast 29.1 44.3 15.2 80.5 36.2
Mali 112.0 124.2 12.2 99.8 -24.4
Niger 235.1 138.0 -97.1 69.3 -68.7
Nigeria 1402.8 607.8 -795.0 489.5 -118.3
Senegal 926.9 875.6 -51.3 712.9 162.7
Togo 15.8 20.0 4.2 28.5 8.5
East Africa 102.6 189.6 87.0 165.3 -24.6
Burundi 7.0 25.0 18.0 57.8 32.8
Egypt 41.5 32.6 -8.9 31.7 -0.9
1.1.2 Area harvested

The average area of groundnut harvested varied considerably from country
to country and from continent to continent between 1960 and 1989 (Tables
1.3 and 1.4). In some countries, this area declined following the use of
new, higher yielding cultivars which resulted in maintained or increased
production on a reduced area. The allocation of land to groundnut culti-
vation is influenced by changes in relative market price and agricultural
policies. In some countries, such as Nigeria, a rapidly growing popUlation
increased the demand for food and this was an important reason for
withdrawing land from groundnut production. In other countries, includ-
ing major exporters such as Argentina, South Africa and Brazil, producers
TABLE 1.1 cant.
Change Change
Country 1960-1969 1970-1979 1970sl1960s 1980-1989 1980s/1970s
Sudan 263.2 621.2 358.0 565.3 -55.9

Tanzania 21.3 58.9 37.6 57.6 -1.3
Uganda 179.9 210.5 30.6 114.3 -96.2
Central Africa 179.2 200.8 21.6 265.3 64.5
Malawi 170.6 158.9 -11.7 176.0 17.1
Zaire 187.7 242.6 54.9 354.5 111.9
Southern Africa 112.6 144.1 31.5 94.5 -49.6
Madagascar 33.6 44.0 10.4 33.2 -10.82
Mozambique 66.8 113.4 46.6 69.0 -44.4
South Africa 251.1 301.2 50.1 179.2 -122.0
Zimbabwe 99.0 117.6 18.6 96.6 -21.0
Asia
South-west Asia 2509.3 2926.5 417.2 3118.1 191.6
India 4951.1 5782.8 831.7 6168.8 386.0
Pakistan 67.4 70.2 2.8 67.3 -2.9
South-east Asia 608.7 669.7 61.0 1221.9 552.2
Indonesia 375.4 504.4 129.0 780.7 276.3
Myanmar 358.8 416.3 57.5 566.7 150.4
Thailand 132.9 210.4 77.5 151.6 -58.8
Vietnam NA 88.2 NA 151.4 63.2
East Asia 289.0 377.0 88.0 412.6 35.6
China 2185.6 2488.7 303.1 4733.9 2245.2
Japan 135.7 87.8 -47.9 50.4 -37.4
Korea 3.5 7.8 4.3 19.4 11.6
Taiwan 110.1 94.4 -15.7 83.9 -10.5
Europe
Turkey 25.4 45.0 24.6 60.2 15.2
reacted to improved production technology, changes in relative price,

agricultural policies and the weather by reallocating land to other crops.
(a) North America

Mexico and the USA are the two major producers on the continent. The
average area harvested in Mexico decreased by 41% during the 1970s,
dropping to a minimum of 25000 ha in 1975, but in the 1980s an initially
slow increase in the area harvested accelerated and the average for the
decade exceeded that of the 1970s by 27%.
The USA harvested, on average, a fairly stable area between 1960 and
1989. The government groundnut programme limited the extension of this
TABLE 1.2 Production tonnage increases and decreases in selected countries
(based on FAG and USDA statistics)
Average 1960s: 1970s 1970s: 1980s

change (%)
Incrcases Decreases Increases Decreases
1-10 Pakistan (Brazil) USA (Pakistan)

Cameroon (Ecuador) India (Egypt)
Burkina Faso (Malawi) Sudan (Tanzania)
(Senegal) Burkina Faso (Gambia)
11-25 Argentina (Taiwan) Mexico (Venezuela)
Paraguay (Egypt) Benin (Ghana)
Myanmar Malawi (Mali)
China (Nigeria)
India (Zimbabwe)
Ghana (Senegal)
Mali (Madagascar)
Uganda (Taiwan)
Zimbabwe
South Africa
Gambia
26-50 Indonesia (Mexico) Colombia (Argentina)
Zaire (Uruguay) Myanmar (Thailand)
Madagascar (Japan) Zaire (Japan)
Togo (Niger) Togo (Cameroon)
(Uganda)
(Mozambique)
(South Africa)
51 and more USA (Nigeria) Paraguay (Brazil)
Venezuela Indonesia (Uruguay)
Thailand Vietnam (Niger)
Sudan China
Ivory Coast Ivory Coast
Benin Burundi
Burundi
Mozambique
Tanzania
area; however, an expansion of the groundnut export market and changes

in the government programme led to a slight increase in area harvested
over time. In the 1980s, it exceeded the 1960s area by 4%.
(b) South America

The area harvested in Argentina varied widely between 1960 and 1989: it
expanded to a record 428000 ha in 1978 and the average area during the
1970s was 19% greater than in the 1960s. However, the introduction of
improved cultivars led to a 39% decrease in that area between 1980 and
1989. In Brazil, the average area harvested decreased by 17% in the 1970s
and continued to decrease steadily in the 1980s. In 1990, Brazilian farmers
planted only 89 000 ha with groundnut, an area 87% less than the record
694000 ha in 1967.
Paraguay expanded its area under groundnut production in the mid
1960s. During the late 1960s and the early 1970s the area figure stagnated
and in some years declined but it increased again between the mid 1970s
and 1990 so that between the 1960s and the 1980s it had grown by 94%.
Several other South American countries cultivate groundnut - for example
Uruguay and Ecuador - but average area harvested has remained below
20000 ha.
(c) Africa
The area harvested in Africa increased during the 1970s but the average in
the 1980s was 5% less than in the 1960s. The land allocation within
individual countries, however, varied in response to each country's specific
demands and policies.
In western Africa, area harvested decreased generally between 1960 and
1990. The area harvested in Senegal remains the largest in Africa, despite
decreasing by 15% during the 1980s. Nigeria assigned the second largest
area to groundnut production in Africa during the three decades but it now
ranks third and the average area harvested in the 1980s was only 2% larger
than its 1960s average. Cameroon emerged as the country with the third
largest area harvested in West Africa: it increased by 90% between the
1960s and the 1980s. In Niger the area harvested decreased by 31%
between the 1960s and the 1970s and by 44% between the 1970s and the
1980s. Mali recorded a decrease of 24% in the period 1960-1989, with an
especially large drop in the 1970s.
In central Africa, the average area harvested increased over time be-
cause of an expansion in groundnut production. Zaire harvested an aver-
age area 77% greater in the 1980s than in the 1960s.
In eastern Africa, the area harvested increased in the 1970s and
remained stable in the 1980s. In the Sudan, the average became the second
largest in Africa in the 1980s, exceeding Nigeria'S area by 21 % . Uganda
decreased its area during the 1970s by 22% compared with the 1960s and
by another 32% during the 1980s. Tanzania increased its average area
harvested by 117% between the 1960s and the 1980s. In Egypt there was a
decrease of 29% during the period 1960-1989.
In southern Africa, the average area harvested in the 1980s was smaller
than in the 1960s. The Republic of South Africa devoted a steadily
decreasing land area to groundnut production. There was no growth in the
TABLE 1.3 A verage area of groundnuts harvested in selected countries
(1000 hectares)
Change Change
Country 1960-1969 1970-1979 1970s/1960s 1980-1989 1980s/1970s
North America
Mexico 69 41 -28 52 11
USA 576 605 29 597 -8
South America
Argentina 284 339 55 173 -166
Brazil 482 399 -83 186 -213
Ecuador 11 12 1 10 -2
Colombia NA 2 NA 3 1
Paraguay 17 20 3 33 13
Uruguay 6 3 -3 2 -1
Venezuela 3 16 13 11 -5
Africa
West Africa 264 291 27 245 -46
Benin 67 74 7 81 7
Burkina Faso 226 173 -53 203 30
Cameroon 166 199 33 315 116
Ghana 44 99 55 105 6
Gambia 145 148 3 99 -49
Ivory Coast 50 51 1 87 36
Mali 161 129 -32 123 -6
Niger 342 235 -107 132 -103
Nigeria 599 953 354 609 -344
Senegal 1063 1106 43 906 -200
Togo 40 31 -9 39 8
East Africa 129 217 88 208 -9
Burundi 9 18 9 50 32
Egypt 21 17 -4 15 -2
average area harvested in Zimbabwe between the 1960s and the 1970s but
it expanded by 9% in the 1980s.
(d) Asia
Asian countries reported the world's largest area harvested of groundnut.
For the continent as a whole, the rapid expansion in average area har-
vested during the 1980s surpassed the 1960s area by 47%. The area in India
is larger than that of any other country and is increasing slowly: it averages
more than three times that of China.
In south-east Asia, the area harvested is substantial. Myanmar is the
TABLE 1.3 cant.
Change Change
Country 1960-1969 1970-1979 1970s/1960s 1980-1989 1980sl1970s
Sudan 320 769 449 739 -30

Tanzania 47 86 39 102 18
Uganda 250 196 -54 133 -63
Central Africa 258 254 -4 374 120
Malawi 239 232 -7 258 26
Zaire 277 276 -1 489 213
Southern Africa 160 176 16 138 -38
Madagascar 40 60 20 35 -25
Mozambique 109 200 91 107 -93
South Africa 319 273 -46 223 -50
Zimbabwe 170 170 0 186 16
Asia
South-west Asia 3506 3604 98 3664 60
India 6967 7159 192 7270 111
Pakistan 45 48 3 57 9
South-east Asia 522 554 32 696 142
Indonesia 380 421 41 579 158
Myanmar 583 651 68 595 -56
Thailand 97 126 29 121 -5
Vietnam NA 84 NA 121 37
East Asia 353 321 -32 354 33
China 1919 2092 173 2647 555
Japan 63 45 -18 28 -17
Korea 4 8 4 15 7
Taiwan 100 69 -31 93 24
Europe
Turkey 11 20 9 24 4
Note Reported figures have been rounded to the nearest thousand.
third largest producer of groundnuts in Asia and its area harvested

increased by 12% between the 1960s and the 1970s, but in the 1980s it was
only 2% above the level of the 1960s. Indonesia increased its average
area harvested by 52% between the 1960s and the 1980s and now ranks
seventh in the world. The increase in Vietnam between the 1970s and the
1980s was an impressive 44%. Thailand expanded its area harvested during
the 1970s but it decreased slightly during the 1980s.
East Asia includes countries with an expanding groundnut sector -
especially China - and countries where groundnut production becomes less
important, such as Japan. China rapidly expanded its area harvested
between 1960 and 1989: the average increased by 9% between the 1960s
TABLE 1.4 Increases and decreases in area of groundnut harvested in selected
countries
Average 1960s:1970s 1970s:1980s

change (%)
Increases Decreases Increases Decreases
1-10 USA (Malawi) India (USA)

Ecuador Guyana (Myanmar)
Pakistan Zimbabwe (Thailand)
China Benin (Sudan)
India (Mali)
Ivory Coast
Gambia
Senegal
11-25 Argentina (Brazil) Pakistan (Ecuador)
Paraguay (Mali) Indonesia (Mozambique)
Myanmar (Uganda) Burkina Faso (South Africa)
Indonesia (Egypt) Malawi (Senegal)
Cameroon (Burkina Faso) Tanzania (Egypt)
Benin (Togo)
(South Africa)
26-50 (Mexico) Mexico (Argentina)
(Uruguay) Colombia (Uruguay)
(Taiwan) Taiwan (Venezuela)
51 and more Venezuela Paraguay (Brazil)
Sudan Vietnam
Ghana Cameroon
Mozambique Ivory Coast
Tanzania Zaire
and the 1970s and by 27% in the 1980s. Japan consistently planted a
reduced area: the average in the 1980s was 56% smaller than in the 1960s.
Korea is a small producer but has been steadily expanding its area under
groundnut cultivation.
(e) Europe
Areas harvested in individual European countries remained below
10 000 ha. Greece, Spain and Italy devoted 3000-5000 ha to groundnut
production. In the early 1960s, Yugoslavia grew 21000 ha of the crop but
the area decreased towards the end of the decade.
Turkey consistently increased its groundnut area: the average increased
by 82% in the 1970s and the expansion continued in the 1980s.
1.1.3 Groundnut yield

Significant changes in groundnut yield occurred between 1960 and 1989
(Tables 1.5 and 1.6). Increased yields, which reflect the use of improved
technology and management, are often necessary to increase exports.
Adoption of improved cultivars adapted to local growing conditions led
to relative increases in yields in China and Argentina during the 1980s.
Increased yields in many other countries signal the potential of cultivars
outside the traditional groundnut-producing countries, though smaller
yield gains have been reported in countries located in climatic zones which
are drastically different from those of the leading groundnut-producing
regions.
(a) North America

Groundnut yields in Mexico and the USA reflect differences in quality of
cuItivars and production technology. Compared with the 1960s, the aver-
age yield in Mexico increased by 13% in the 1970s but then decreased by
5% in the 1980s. In contrast, the USA average yield consistently increased,
in relative terms, by 62% between the 1960s and the 1980s.
(b) South America

Between the 1960s and the 1980s, average yield in Brazil increased by
20%. Brazil experienced periods of declining yield in the late 1960s and the
first half of the 1980s, but the average has increased steadily overall.
Changes in yield were more dramatic in Argentina. The average yields
were smaller than in Brazil during the 1960s and 1970s, but larger in the
1980s when Argentina exceeded its 1970s yield by 41 %.
Yield in Paraguay and other South American countries remained con-
siderably lower than in Argentina and Brazil throughout the three decades.
However, the average yield in Paraguay increased by 38% and the rate of
yield increase was especially high during the 1980s.
(c) Africa
A number of groundnut producers in Africa achieved record yields in the
1960s and 1970s but experienced a decline during the 1980s. Among
countries reporting this pattern were Benin, Burkina Faso, Burundi, Mali,
Tanzania, Uganda, Zaire and the Republic of South Africa.
In western Africa, the overall average yields remained stable between
1960 and 1990 but fluctuated in individual countries. The average Gambian
yields have been consistently above 500 kg/ha since 1983. Senegal reported
considerably lower and more variable yields than Gambia. Yields in
Nigeria, a large producer of groundnuts, exceeded its average yield of the
1970s by 32% in the 1980s. In Ghana and Cameroon average yields
TABLE 1.5 Average groundnut yields in selected countries (kg/ha)
Change Change
Country 1960-1969 1970-1979 1970s/1960s 1980-1989 1980sl1970s
North America
Mexico 1329 1497 168 1420 -77
USA 1724 2652 928 2792 140
South America
Argentina 1154 1174 20 1660 486
Brazil 1231 1383 152 1481 98
Ecuador 985 956 -29 925 -31
Colombia NA 1600 NA 1302 -298
Paraguay 822 869 47 1137 -268
Uruguay 735 1058 323 550 -508
Venezuela 987 1235 248 1501 266
Africa
West Africa 743 757 14 752 -5
Benin 379 751 372 694 -57
Burkina Faso 510 651 141 601 -50
Cameroon 716 620 -96 502 -118
Gambia 706 855 149 1165 310
Ghana 1708 761 -947 559 -202
Ivory Coast 596 872 276 916 44
Mali 792 1124 332 856 -268
Niger 724 540 -184 508 -32
Nigeria NA 619 NA 817 198
Senegal 876 781 -95 804 23
Togo 423 756 333 846 90
East Africa 977 1278 301 1092 -186
Burundi 696 1496 800 1073 -423
Egypt 2004 1920 -84 2201 281
declined between 1960 and 1989 - in Ghana by 67% and in Cameroon by

30%. The average yield in the Ivory Coast climbed by 54% over the same
period. In Benin and Burkina Faso the average 1970s yields were higher
than in the 1960s or the 1980s; Niger reached its highest average yield in
the 1960s and Mali in the 1970s.
In eastern Africa average yields vary widely. Egypt's, the highest in
Africa, increased by 10% between the 1960s and the 1980s. Uganda's yield
increased by 72% during the 1970s compared with the 1960s but the
average then decreased by 31 %. Tanzania reported a rapid increase of
average yield during the 1970s but, again, these gains were eroded during
the 1980s and decreased by 37%. In relative terms, average yield in
Mozambique decreased by 62% between 1960 and 1989. The Sudan also
TABLE 1.5 cant.
Change Change
Country 1960-1969 1970-1979 1970s/1960s 1980-1989 1980s/1970s
Sudan 835 817 -18 753 -64

Tanzania 574 910 336 572 -338
Uganda 724 1245 521 860 -385
Central Africa 699 794 95 711 -83
Malawi 715 684 -31 681 -3
Zaire 682 903 221 742 -161
Southern Africa 866 792 -74 678 -114
Madagascar 875 845 -30 960 115
Mozambique 1073 534 -539 413 -121
South Africa 796 1106 310 801 -305
Zimbabwe 718 682 -36 536 -146
Asia
South-west Asia 1050 1139 89 1015 -124
India 712 807 95 841 34
Pakistan 1388 1470 82 1188 -282
South-east Asia 997 1199 202 1136 -63
Indonesia 988 1185 197 1506 321
Myanmar 620 644 24 950 306
Thailand 1382 1671 289 1255 -416
Vietnam NA 1297 NA 833 -464
East Asia 1465 1505 40 1543 38
China 1140 1195 55 1760 565
Japan 2148 1943 -205 1773 -170
Taiwan 1107 1376 269 1092 -284
Europe
Turkey 2332 2289 -43 2324 35
suffered a slow but steady decline of 10% in average yield between the
1960s and the 1980s.
In central Africa, average yields increased but then decreased, leaving
them at a slightly higher level in the 1980s than in the 1960s. In southern
Africa, the average yield declined steadily between 1960 and 1989. The
Republic of South Africa, a major African producer, experienced an
increase in average yield during the 1970s but the average during the 1980s
was 28% lower than in the 1970s. Zimbabwe's yield declined by 21 % in the
1980s compared with the 1960s.
(d) Asia
In south-west Asia, India's average yield increased by 13% between the
1960s and the 1970s and by 4% in the 1980s.
TABLE 1. 6 Yield increases and decreases in selected countries in comparison to
average yields in the period 1960-1969 (based on FAG and USDA statistics)
Average 1960s:1970s 1970s:1980s

change (%)
Increases Decreases Increases Decreases
1-10 Argentina (Japan) USA (Mexico)

Paraguay (Egypt) Brazil (Ecuador)
Myanmar (Sudan) India (Japan)
China (Malawi) Ivory Coast (Benin)
Pakistan (Zimbabwe) Senegal (Burkina Faso)
(Cameroon)
(Niger)
(Sudan)
11-25 Mexico Venezuela (Colombia)
Brazil (Senegal) Egypt (Pakistan)
Venezuela (Niger) Togo (Taiwan)
Taiwan (Thailand)
Thailand (Zaire)
India (Mali)
Indonesia (Zimbabwe)
Gambia (Mozambique)
26-50 Uruguay Argentina (Uruguay)
Burkina Faso (Cameroon) Paraguay (Vietnam)
Ivory Coast (Mozambique) Myanmar (Ghana)
Mali China (South Africa)
Zaire Indonesia (Tanzania)
South Africa Nigeria (Uganda)
Gambia
51 and more USA (Ghana)
Benin (Nigeria)
Tanzania
Uganda
Yields in eastern Asia increased in the 1970s but decreased slightly in the
1980s. However, Myanmar's average yield in the 1980s was 48% higher
than in the 1970s. Yields in Thailand peaked between 1975 and 1977 but
declined below these record levels: the average yield in the 1980s was 9%
lower than in the 1960s.
Yield in Indonesia increased over time. The average increased by 20% in
the 1970s and was 52% higher in the 1980s than in the 1960s. Vietnam is
expanding its groundnut production but its average yield in the 1980s was
36% below that of the 1970s.
In south-east Asia, yields in Taiwain and Japan were on average lower in
Stocks and prices 15
the 1980s than in the 1960s. Yields in China remained stagnant during the
1960s and 1970s but then increased, reaching a record 2036 kg/ha in 1987,
and decreased again in 1989-1990. The average yield in the 1980s was 48%
higher than in the 1970s. The average yield in Korea has been increasing
rapidly but its groundnut sector remains small.
(e) Europe
Competition from other crops and a highly developed agriculture in
Europe contribute to average yields which are among the highest in the
world. The exception is the former USSR but groundnut is produced
largely in non-European regions of the Commonwealth of Independent
States.
Among small European producers, Italy attains high average yields
which frequently exceed 2 000 kg/ha. Spain and Greece have also achieved
high yields but those of Yugoslavia and Bulgaria have been highly variable.
The largest groundnut producer in Europe is Turkey, whose average
yield in the 1960s was 2332 kg/ha. This decreased by 2% in the 1970s and
in the 1980s it was 0.3% below the 1960s level.
1.2 STOCKS AND PRICES
The level of world groundnut stocks influences both supply and prices in
the groundnut trade. A record level of 944 000 MT was achieved in 1976177
as the result of accumulation of large stocks in the USA, India and
Argentina. An increased demand for oil-bearing crops in the preceding
year was probably a major factor in this accumulation. Large domestic
stocks in Argentina and the USA led to large exports in the following
years.
Between 1971172 and 1990/91, the USA consistently maintained the
largest stocks in the world. These ranged from 178000 MT in 1972173 to
646000 MT in 1985/86 but declined in the late 1980s. India recorded its
highest stocks (450000MT) in 1974175, following the world food shortage
of the early 1970s, but the levels have varied widely from year to year.
Since 1986/87, India has reported no groundnut in stock. Argentina
reported large stocks between 1975176 and 1977178 but levels decreased
during the 1980s.
The combined stocks of other countries have decreased from a record
level of 615000 MT in 1974175; during the 1980s, they exceeded
400000 MT in 1982/83 only. In 1990/91, the groundnut inventory was
169000 MT in these other countries of the world - excluding the European
Community (EC) countries, which maintained stocks ranging from none to
58000 MT between 1971172 and 1990/91.
Stock levels and price influence domestic consumption and the level of
exports. Changes in price in major producing and exporting countries
change export levels. During the 1960s, groundnut prices were the lowest
in African countries: pricing was particularly competitive in Senegal,
Gambia and the Sudan and relatively low in the Republic of South Africa,
Malaysia and Nigeria, while prices in China, Argentina and the USA were
amongst the world's highest.
Prices remained low in African countries in the early 1970s but gradually
increased until those in the Sudan and the Republic of South Africa were
among the highest in the world by the late 1970s. Between the late 1970s
and the early 1980s, the USA, Argentina and China were more competi-
tive in relation to African exporters.
In the USA, prices were supported under a government programme.
The early 1980s brought a change in the programme through the introduc-
tion of a two-tier price system. Groundnut sold at a higher price for quota;
production above the quota level, selling at a lower price, was called
'additionals'. The price of additionals responded to the expected prices of
groundnut oil, meal, other vegetable oils and world groundnut demand.
The two-tier price system lowered the price of exported groundnut. The
price on the world market became less influenced by the US government
programme.
Table 1.7 shows export prices for major exporting countries from 1960 to
1987. The USA and Argentina became more competitive in the 1970s, as
did China in the late 1970s when it began its large-scale exports. During the
1980s, the prices of many exporters became less attractive to buyers, given
the quality and the intended use.
International groundnut price provides information to producers,
brokers, and consumers. The price for each type of groundnut reflects
supply and demand conditions and the quality attributes of traded
cultivars.
The supply situation changes frequently because harvests in the northern
and southern hemispheres occur at different times of the year. A major
factor that influences the supply is the weather. Weather-related supply
changes cause price variability and alter quantities exported by specific
countries. Quality also influences price, but crop quality information is less
readily available. Groundnut quality is determined during trading and
processing.
Prices in different parts of the world interact with each other, reflecting
the many uses of groundnut. Groundnuts from different sources are
reasonable substitutes, especially for a similar use. Some specialized end
users, however, prefer groundnut with specific quality attributes and pay
higher prices to satisfy their needs.
Timely information concerning current and expected price on the inter-
national market affects planting decisions, stocks and quantities of applied
inputs. A European-based company publishes prices of the three major
groundnut exporters: the USA, China, and Argentina. Reports include
International trade 17
TABLE 1.7 Groundnut export prices for major exporters (US dollars per ton)
(based on FAG statistics)
Year US CN AG SD SAF GB ID BZ THL SEN VT MAL
1960 235 247a 257 192 195 185 242 NA 188 173 NA NA
1961 255 247a NA 186 188 166 276 220 183 172 NA NA
1962 318 233 a 257 163 188 160 254 185 184 173 NA NA
1963 239 235 a NA 159 184 150 233 168 198 173 NA NA
1964 228 234 a 225 173 186 154 273 190 197 173 NA 198
1965 251 232a 200 155 219 199 205 222 202 172 NA 288
1966 251 221 a 174 210 208 163 96 251 196 175 NA 243
1967 218 233 a 211 178 183 154 208 230 305 174 NA 185
1968 230 233 a NA 161 191 133 213 228 295 137 NA 185
1969 358 269a 333 230 197 185 280 222 309 165 NA 196
1970 382 275 a 393 246 197 194 293 229 307 188 NA 226 a
1971 292 297a 296 229 214 233 287 247 317 197 NA 244
1972 261 319a 303 242 207 223 278 244 401 269 NA 251
1973 435 447 3 481 275 275 267 470 359 470 387 NA 268
1974 600 640 464 548 334 463 507 557 633 573 NA 300
1975 622 680 516 482 332 493 552 606 a 65Y 530 NA 291
1976 685 560 507 396 297 453 753 634 682 380 242 473
1977 693 642 839 577 397 550 969 849 723 477 290 531
1978 733 710 1000 641 418 a 529 1408 853 702 602 429 815
1979 741 701 526 715 581 569 743 857 864 948 614 798
1980 731 728 628 518 947 409 805 793 1380 807 620 767
1981 943 1047 942 1037 792 670 1192 1241 1195 1116 625 693
1982 864 702 559 385 614 374 889 863 835 408 603 612
1983 784 590 440 796 692 389 912 905 1042 435 589 625
1984 762 713 700 833 750 535 947 1025 760 450 652 658
1985 676 679 450 763 644 490 652 845 619 619 553 382
1986 727 588 502 905 619 637 571 893 566 527 419 453
1987 787 686 435 612 671 606 762 930 479 412 385 323
aEstimate
NA = not available
US = United States, CN = China, AG = Argentina, SD = Sudan, SAF =
Republic of South Africa, GB = Gambia, ID = Indonesia, BZ = Brazil, THL =
Thailand, SEN = Senegal, VT = Vietnam, MAL = Malawi.
prices for the US Runner 40/50s, Argentine Runners, and Chinese Hsujis.
Quarterly price quotations allow traders and producers to adjust expec-
tations and make appropriate decisions.
1.3 INTERNATIONAL TRADE
The strong world demand for groundnut oil on the one hand and ground-
nut products on the other determines the economic significance of the
crop. This dichotomy of demand has led to the evolution of two markets.
Groundnuts traded as an oil-bearing crop compete with other oil-bearing
crops such as palm oil, soybean, rapeseed, etc. - groundnuts as a source of
oil are important in many lcss developed countries. Used as a snack,
groundnuts compete with other snack foods "for example, tree nuts. In the
form of butter, groundnuts compete with other spreads. Groundnut snacks
and peanut butter are consumed largely in developed industrial economies.
Quality characteristics of groundnuts as snack foods or as kernels for oil
reflect the two distinct uses of groundnut that influence the volume traded,
price, and quality attributes. Groundnuts intended for different uses follow
separate geographical trade patterns and use specialized marketing chan-
nels. The needs of end-users communicated through market price influ-
ence the selection of cultivars and levels of production. A separate market
exists for groundnut meal and cake (by-products from groundnuts crushed
for oil), which compete with other feed protein supplements and are
purchased primarily by countries with large commercial livestock
production.
1.3.1 International trade in groundnuts

The volume of world trade increased slowly during the period 1971172-
1990/91. During the 1970s, the volume traded on the world market ranged
between 950000 MT and 1200000 MT - except in the 1975176 season,
which was characterized by severe drought, especially in Europe. This
drought led to increased demand for imported oil-bearing crops. During
the 1980s, the volume traded edged upwards, although at an erratic
pace, and ranged from 1000000 MT to 1300000 MT. In the years 1985/86
and 1990/91 the volume traded was 1365000 MT and 1339000 MT,
respectively.
The geographical composition of exporters changed between 1971172
and 1990/91. The USA exported the largest volume during the period
1971172-1974175. After a short crop in 1975, Europe looked for alternative
sources of oil and meal and imported groundnuts as a substitute for other
oil-bearing commodities which, in normal crop years, were also less expen-
sive. Simultaneously, according to the existing programme in the USA, the
price support for groundnuts increased to compensate for increased pro-
duction costs, resulting in the highest price of any major exporting country.
This encouraged importers to search for other suppliers: the Sudan
exported 293000 MT in 1976; India exported 239000 MT, while Senegal
reported record exports of 173 000 MT. Countries which surpassed the
USA in exports in 1976 were those with lower prices: the price of Sudanese
groundnut was 42% lower and Indian 30% lower.
Before 1980, other major exporters included the Republic of South
Africa, the Sudan and Senegal. Since 1980, both China and Argentina have
International trade 19
surpassed the Republic of South Africa and the Sudan in export volumes.
A short 1980 crop in the USA led to difficulties in meeting importer's
demands and the USA itself imported 82000 MT in 1980/81, leaving net
exports of only 46000 MT.
The shortage in the USA groundnut supply coincided with changes in
economic and agricultural policies in China. China's groundnut exports
increased following the change in regulating domestic agricultural pro-
duction and the implementation of economic incentives for farmers. China
exported 305000 MT in 1980/81; the figure declined between 1981/82 and
1984/85 but China has since exported the largest quantity of groundnut in
the world, except for 1986/87 and 1989/90.
Argentina had a history of groundnut exports dating back to World War
II. Despite the deteriorating terms of trade during the period 1935-1946,
Argentina expanded its exports in 1940-1945 (Mundlak et al.,1989). For
years its groundnut exports were negligible but they were increased follow-
ing record production in 1976177 and 1978179 (a result of increased yields
and a near-record area harvested). The need to generate foreign exchange
caused a steady increase in the volume exported and the quality of
Argentine groundnut satisfied buyers' requirements. Between 1985/86 and
1989/90, volume exceeded 100000 MT; in 1990/91, exports from Argentina
reached a record 215000 MT. Furthermore, domestic economic policies
maintained a competitive edge and Argentina's groundnut price between
1984 and 1990 was the lowest among major exporting countries.
The USA, China and Argentina will continue to share the bulk of the
world export market in the foreseeable future. Argentina can easily
expand the area harvested, while the USA and China can alter allocation
of land to groundnut production, depending on domestic needs and oppor-
tunity costs.
Changes in quality and consumption have influenced shifts in origins of
groundnut exports. Groundnuts, intended for use as a source of edible oil,
have become over time a food item eaten as kernels or paste. This change
called for different quality attributes. Crushing qualities remained import-
ant to oil producers but appearance, flavour and texture became more
important for snack foods and peanut butter.
The composition of the import market remained stable but a few
countries, such as Indonesia, Korea, and Malaysia, increased imports after
1979. Major importers included Canada, western European countries (12
EC nations and Switzerland, Sweden and Norway), Japan and the former
USSR. Total imports by Canada and Japan remained largely unchanged,
while the USSR imports increased steadily. Since reaching a peak in 1976,
western European countries have reduced total imports to about 50% of
1976 levels.
The three major competitors in the western European market are the
USA, Argentina and China. The USA market share in western Europe
increased from 14% in 1973 to 37% in 1986, with a peak of 54% in 1979.
Western European countries absorbed 60% of the USA exports.
Argentina and China increased their shares from 7% and 8% in 1982 to
20% and 21 % in 1986, respectively. The Republic of South Africa has also
been a significant exporter to the western European market but has lost
more than half of its market share.
China and the USA have shared equally about 85% of the market in
Japan, with some annual variation in quantities exported. The USA
shipped about 10% of its groundnut exports to Japan. Argentina also
exported small quantities to Japan. The USA dominated the Canadian
market for years: from 1973 to 1986, Canada alone bought more than 20%
of US groundnut exports each year, except in 1980. Chinese exports are
increasing steadily and, while the market share of Chinese groundnut in
Canada accounted for only 5% in 1982, it increased to 11% in 1984 and
exceeded 15% in 1986. The major sources for Indonesia are China and
Vietnam, with an increasing importance for the latter. The opening
of Indonesia's market to Vietnam groundnuts coincided with improved
political relationships between the two countries.
A relatively strong dependence on exports characterizes the USA's
position in the world market. The USA is not the largest groundnut
producer in the world but is consistently the largest exporter. The position
of Argentina is even more sensitive to world markets because the ground-
nut is largely an export commodity: the domestic market is small and
Argentina exports groundnut to earn foreign currency. China, the second
largest producer, uses most of its crop for domestic consumption; exports
represent only a fraction of its total production and can be expanded.
1.3.2 International trade in groundnut oil

Groundnut oil is an important vegetable oil in some regions where other
oil-bearing crops are not grown; in other regions, it is a substitute for other
vegetable oils. In some cultures, consumption habits call specifically for the
use of groundnut oil in cooking.
World exports of groundnut oil increased sharply between 1960 and
1968, reaching 510 000 MT, but the trade decreased in the late 1960s.
African exports dominated the world oil market and the varying volume of
ground nut oil on that market resulted from the variable supply coming
from Nigeria and Senegal. Senegal was the largest exporter in the 1960s,
followed by Nigeria and then Argentina.
During the 1970s, the most significant change was a steady decline of
Nigerian oil shipments. Towards the end of the decade Nigeria became a
net importer of groundnut oil. Senegal remained the largest ground nut oil
exporter but at highly variable rates during the 1970s. Brazil and China
emerged as new oil exporters in the 1970s: Brazil slowly expanded its
exports and maintained them through the decade, while China's oil exports
were especially high during the first half of the 1970s and at the end of the
Concluding comments 21
decade. Argentina and the Republic of South Africa continued to export
oil during the 1970s.
In the 1980s exports from Senegal, Argentina and Brazil decreased and
the Sudan began to export significant quantities of groundnut oil.
Major buyers include the EC members (purchasing oil from Argentina
and Africa) and the countries of Asia where the groundnut oil trade is
largely intra-continental.
1.3.3 International trade in groundnut cake and meal

The trade in groundnut cake and meal followed a different pattern from
the trade in shelled nuts or oil. European countries were the major buyers
of meal in the first half of the 1960s; Japan and the Soviet Union also
purchased significant quantities of cake and meal.
In the first half of the 1960s, India was the largest cake and meal
supplier, its share frequently exceeding 50% of the total world cake and
meal exports. Argentina and Senegal were other major exporters. In the
second half of the decade, India and Senegal continued to export large
quantities of groundnut cake and meal but Argentina reduced its exports
while Brazil rapidly increased its share of the world cake and meal market.
During the 1970s, India and Senegal continued to dominate cake and
meal exports; the importance of exports from Argentina decreased
and the Sudan's exports slowly expanded. After peaking in the early
1970s, Brazilian exports declined. Myanmar exported fairly large quanti-
ties of cake and meal in the 1960s but its share decreased by the mid
1970s.
The international trade decreased during the second half of the 1970s
and the presence of aflatoxin in cake exported from Senegal may have
contributed to a decrease in exports to Europe. A decrease in trade did not
change the relative importance of India's and Senegal's cake and meal
exports. The Sudan emerged as a large cake and meal supplier towards the
end of the decade; Myanmar decreased its exports while exports from
Argentina and Brazil were highly variable. Europe remained the primary
destination for cake and meal exports.
In the 1980s, the trade continued at the 1970s level, according to the
geographical pattern established in earlier decades. The exception was the
year 1985, when exports reached a record low level of 385 000 MT.
1.4 CONCLUDING COMMENTS
This overview of the world groundnut yields, area harvested and pro-
duction indicates considerable regional differences. Groundnuts are culti-
vated in many countries but their relative economic importance varies. As
in the case of many other cultures, production is highly developed in
countries with well-functioning agriculture. Differences in applied
technology and management are major factors contributing to regional
differences in yield and production. Marketing infrastructure (including
transportation, storage, quality control, advertising and promotion) is an
important factor influencing consumption of groundnut and groundnut
products.
The quality-price relationship determined by end users is likely to
increase in importance, and is influenced by both biological and economic
factors. Quality attributes preferred by buyers can be met by cultivar
development. The increasing importance of health concerns resulted in the
expansion of canola production which has become the largest oil crop in
the world (Fletcher et al., 1992). Re-examination of peanut oil properties is
a desirable long-term goal. Protecting quality after harvest and the timely
supply of adequate volume are essential for the smooth operation of
regional and international markets.
Opportunities and constraints on the world market will be generated
primarily by variable production in different countries and regions.
Droughts, sudden disease outbreaks or insect attacks can occur with little
warning, limiting time for adjusting market strategy. Canada, Japan and
western Europe will remain the largest buyers. Large but variable quan-
tities have been exported to Indonesia. Expanding markets include
countries with well-developed industry and service sectors, such as those of
the Pacific Rim. New markets include the central and east European
countries where economic reforms are being implemented. Although
groundnut products can become easily available, they remain expensive.
The former USSR represents a special case. It was steadily increasing
its imports, their volume being comparable in size to Canada's.
political and economic changes and the emergence of new independent
states will bring changes in lifestyle and consumption which may help
exports. Some of the new states may increase their own production after
the system of central planning disintegrates and privatization of the econ-
omy progresses.
ACKNOWLEDGEMENTS
The author would like to thank D. Cummins, S. M. Fletcher and J. C.

Purcell for their helpful comments on earJi.er drafts of this chapter.
REFERENCES
Fletcher, S.M., Zhang, P. and Carley, D.H. (1992) Peanuts: Production, Utilization and
Trade in the 1980s. Department of Agricultural and Applied Economics, College of
Agriculture, University of Georgia, FS-91-32.
References 23
Food and Agricultural Organization Production Yearbook, United Nations Organization,
various issues.
Food and Agricultural Organization Trade Yearbook, United Nations Organization, various
issues.
Mundlak, Y., Cavallo, D. and Domenech, R. (1989) Agriculture and economic growth in
Argentina, /9/3-/984, IFPRI, Research Report 76.
US Department of Agriculture World Oilseed Situation and Market Highlights, Foreign
Agriculture Circular, various issues.
CHAPTER 2
The origin and history

of the groundnut
R.O. Hammons
2.1 INTRODUCTION
Groundnut is a native New World crop. Early explorers found it culti-

vated extensively in both Mesoamerica and South America. Remnant
pericarp (fruit hull) tissue recovered from archaeological sites in Peru
dates its purposeful agricultural use there at approximately 3900-3750
years before the present (YBP). No one is certain how much earlier
domestication occurred but it probably first took place in the valleys of
the Parana and Paraguay river systems in the Gran Chaco area of South
America.
Today, groundnut is an important oil, food and forage crop generally
distributed in tropical, subtropical and warm temperate zones. The exact
origin of the principal cultigen, Arachis hypogaea L., remains a subject of
scientific inquiry. Early Spanish, Portuguese, Dutch, German and other
explorers found Indians cultivating the crop on many islands in the
Antilles, on the northeast and east coasts of Brazil, in all the warm
regions of the Rio de la Plata basin (Paraguay, Bolivia, northern
Argentina, extreme southwest Brazil), extensively in Peru and sparsely in
Mexico.
This chapter relates the history of the crop as revealed in art, artifacts
and archaeological remains from prehistoric sites; it traces early post-
Columbian accounts of the plant or fruit in contemporary natural historical
narratives of the sixteenth and seventeenth centuries and reviews current
information concerning the crop's centre of origin and geographic disper-
sion. It also lists linguistic affinities in common names and notes the impact
of the European vegetable oil industry on production expansion in West
Africa.
The Groundnut Crop: A scientific basis for improvement. Edited by J. Smartt. Published in
1994 by Chapman & Hall, London ISBN 0 412 408201.
Prehistoric ground nut agriculture 25
2.2 PREHISTORIC GROUND NUT AGRICULTURE
Peruvian civilization apparently began along the eastern slopes of the

Andes near the tropical Amazonian lowlands, where many wild relatives of
domesticated crops occur (Burger, 1989). However, archaeological evalu-
ation of agricultural sites in that area is difficult because of climatic
conditions. The coast of Peru at about 11° S is an extremely dry desert due
to the cold off-shore ocean currents and no natural vegetation exists
(Cohen, 1977). Along these shores are the oldest known examples of
monumental architecture in the New World. The native peoples reshaped
their environment to create distinctive Andean agricultural systems. About
4200 YBP the coastal society began building small scale irrigation systems
(Burger, 1989). By 3200 YBP the demographic, technological and socio-
economic foundations for central Andean civilization were in place.
The finding of well-preserved groundnut fruits in terracotta jars in the
prehistoric burial sites at Ancon on the Peruvian coast north of Lima
conclusively demonstrated the New World origin of the cultivated crop
(Squier, 1877). Specimens recovered there date to the early Ancon culture,
approximately 2740-2490 YBP (Towle, 1961).
Further north near Trujillo, archaeologists found funerary vases decor-
ated with replicas of groundnut pods sculptured in relief, some of which
used moulds made from actual pods. An earthenware pan recovered from
a grave at the fishing village of Chimbote, Peru, had groundnut fruits
painted upon the handle.
From about 1900 to 1300 YBP the agricultural peoples of the Mochica
culture grew irrigated crops such as maize, beans, squash, peppers, pota-
toes, manioc and groundnut amid the searing desert of coastal Peru. The
Moche empire extended along a 136 km swath of coastal Peru. Recently,
near the village of Sipan, 7.5 km west of Pampa Grande in the heart of the
Lambayesque valley, Walter Alva (1988) unearthed the richest docu-
mented burial ground of a pre-Hispanic ruler, finding treasures of unprece-
dented artistic magnificence. From above the 'royal' tomb they recovered
from clandestine looters a pair of gold groundnut pods three times larger
than natural size. The artisans faithfully reproduced the ridges and inden-
tations that are present on one major cultivar grown in the area even today.
Two unusual necklaces exhibited quality in attention to detail and care in
execution: half of each necklace was of silver and the other half gold,
fashioned into groundnut pods. A third necklace displayed a gold ground-
nut pod but its alternating lapis and gold beads reveal an extensive Moche
trade network, with gold originating in the eastern Andes and the lapis
from Chile.
The best dates yet established for the occurrence of groundnut in Peru
are Bird's (1948) finding of its remains at Huaca Prieta near latitude 8° S on
the coastline of the Chicama valley. Bird placed the appearance of ground-
nut there prior to maize and probably contemporaneous with warty
26 The origin and history of the groundnut
squash. The precedence of groundnut to maize here is unique for Peruvian
sites. Neither groundnut nor warty squash plant remains appeared in the
preceramic refuse, suggesting their introduction concurrent with the first
pottery. Carbon dating of the pottery, and thus for groundnut, is for the
beginning of the ceramic period and ranges from 3500 to 3200 YBP.
However, Cohen (1977) reports groundnut remains from the Gaviota
Complex (preceramic period 6) in the Anc6n-Chill6n region of the coast of
Peru with a possible age of 3750-3900 YBP. Lima bean, jack bean, sweet
potato, peppers and squashes occurred with groundnut in this period, but
not maize or the common bean that later became staples of Peruvian
agriculture.
The usual cultivar of A. hypogaea found in the coastal sites of Peru has
the long, slender, ribbed pod and sharp, recurved beak typical of subspe-
cies hypogaea var. hirsuta (Krapovickas, 1968). Its similarity in external
morphology, pod size and seed number to a major phenotype sold today in
markets of coastal Peru reflects its antiquity in cultivation in ancient Peru.
Towle (1961) dates remains of a smaller podded type that she recovered
from a site at Supe, Peru, to the Early Anc6n period. The modest retic-
ulation and lack of dorsal humps on its pods suggest selection under
domestication. The presence of groundnut, manioc and chilli pepper -
domesticated east of the Andes - in early ceramic cultures to the west
shows prolonged and effective contact between residents of the arid coast
of Peru and those of the Gran Pantanal region.
The discovery of ethnobotanical samples of cultivated groundnut in a
Coxcatlan cave in the Tehuacan valley of Mexico established groundnut as
a cultigen on both American continents many centuries before their dis-
covery by the Europeans. These date to about 1900 YBP. Although present
in later phases (1200--450 YBP), they were never abundant (MacNeish,
1965; Smith, 1967). The surviving Aztec codices lack known illustrations of
the ground nut. Current evidence suggests that ground nut was an intro-
duced and unimportant crop in Mexico.
Phytomorphic representations of groundnut have yet to be found in
Brazil, Bolivia or the Antilles, where the climate is less favourable for the
preservation of archaeological plant remains. Thus, for the present, the
deductions drawn from archaeological data must be supplemented and
extended by evidence from natural historians and other disciplines.
2.3 POST -COLUMBIAN HISTORICAL NARRATIVES
2.3.1 Sixteenth century natural histories

The first written notice of the groundnut in the chronicles and natural
histories of the sixteenth century is that of EI Capitan Gonzalo Fernandez,
subsequently known as de Oviedo y Valdes, the first historian of the New
Post-Columbian historical narratives 27
World. Ferdinand and Isabella sent Oviedo to Santo Domingo in 1513,
where he soon became governor of Hispaniola (now Haiti/Dominican
Republic) and royal historiographer of the Indies. In 1525 he sent Emperor
Charles V his Sumario Historia, printed in Toledo two years later (Oviedo,
1527), and in 1535 he began publishing his His to ria General de las Indias
(Oviedo, 1535, 1547), a task finally completed three centuries later by the
Spanish Royal Academy of History (Oviedo, 1851).
Oviedo (1527) describes groundnut (manzl as a very abundant, ordinary
food crop sown and harvested in the gardens and fields of the Indians on
Hispaniola and other islands. Its fruit was about the size of a pine nut with
the shell. The Indians considered groundnut a healthy food and consump-
tion was high.
Three centuries later the completed Cronica reported that the fruit grew
underground, that pulling upon the branches uprooted the plant for
harvest and that the seeds eaten raw or roasted were very tasty (Oviedo,
1851). These statements are not in the several sixteenth century editions
(Hammons, 1982).
Although the specific geographic area where farmers domesticated
groundnut is unquestionably South America, Oviedo's 1527 narrative also
established distribution and adaptation of the crop as far as the West
Indies, and its widespread and ordinary usage there before the discovery of
the Americas by Europeans.
Ulrich Schmidt of Straubing, a German adventurer who spent 20 years in
the Rio de la Plata basin, subsequently wrote a historical account of the
conquests by the emperor's forces there. Schmidt (1567) encountered
groundnut in 1542 when his expedition up the Rio Paraguay from
Asunci6n met the Cheriguanos (or Guaycunis) Indians who bartered
maize, manioc and groundnut with the soldiers. Later, in the area
inhabited by the Mepenes (or Abipones) he recorded a flourishing agri-
culture with maize, manioc, potatoes, sweet potatoes and groundnut in
abundant supply. I
The first delibrate introductions of groundnut to Europe probably went

unrecorded. The New World presented a bountiful wealth of domesticated
seed, root and tree crops, including maize, the amaranths, four species of
bean, warty and summer squash, groundnut, pumpkin, gourds, tomato,
chilli pepper, potato, manioc, sweet potato, the New World yam, arrow-
root, cacao (Theobroma), papaya, pineapple, the cultivated tobaccos and
cotton, inter alia. From the time of Columbus' first voyage, explorers
collected useful and exotic American plants for introduction to Europe.
One may speculate on the introduction of groundnut - plentiful in the
Antilles - to Europe early in the sixteenth century. However, extensive
reviews of early literature by writers of groundnut history (Dubard, 1906;
Higgins, 1951; Krapovickas, 1968; Gillier and Silvestre, 1969; Hammons,
1973, 1982) do not document such an occurrence.
The writer who first gave Europe an account of the more useful plants
discovered by the Spanish adventurers in the New World was Dr. Nicholas
Monardes, a physician of Seville, who published a small treatise on medici-
nal plants (Monardes, 1569). Other Spanish editions appeared in 1571 and
1580, and the work passed through many editions in many languages,
including an English version in 1577 by Frampton. Better known than the
original Spanish or the vernacular translations is the excellent Latin version
included in the Exoticorum of Clusius (1605), which spread this knowledge
throughout the world of learned men.
Monardes never visited America but derived his information and speci-
mens from the navigators and explorers who frequently arrived in Seville.
He saw only the nameless underground fruit, which he described as having
neither root nor plant! He wrote that the fruit was:
half a finger round and had a small kernel [sic] that parted into two
halves ... It may be eaten green (raw) or dry, but the best way is to
toast it.
Since the samples came to Monardes from the sandy banks (,coasts') of the
Rio Marafion, he postulated that this area was the only part of the 'Indias'
where the fruit grew. As a cultural note, he observed that both the Indians
and the Spanish in Peru held the groundnut in great esteem.
The second writer to publish a book upon American natural history was
Jean de Lery. A Calvinist missionary with the Huguenot colony founded
by the Chevalier de Villegagnon in 1555 on a small island in the bay of Rio
de Janeiro (now Guanabara bay), Lery remained in Brazil less than five
years. In 1578, he published the account of his voyage to the land of Brazil,
describing groundnut (manobt) as 'a fruit the savages' grew in the soil like
truffles. He wrote that it had a hard pod containing grey-brown seeds with
size and taste similar to the hazelnut. Although Lery said he had eaten the
fruit often, he could not say whether the plant had leaves or seed.
At about the same time as Monardes thought groundnut grew only along
the river Marafion, the Portuguese naturalist Gabriel Soares de Souza
(who lived in Brazil from 1570 to 1587) believed that the plant occurred
only in that country. Soares (1587) described the groundnut plant, its
cultivation and the common native custom of using smoke to cure the crop.
His detailed account of production practices is still of interest: the seed was
planted 'a hand's breadth apart' in February, in humid soil prepared solely
by Indian or halfbreed females who, in May, each harvested what they had
planted. The plant's leaves resembled those of Spanish beans, and the
branches trailed along the ground. Soares mistakenly placed the fruit pods
on the root ends. The pods resembled pine nuts but had a thicker shell and
contained three or four seeds each. The seed, when eaten raw, had the
taste of chickpea. In-shell roasting was the usual method of preparation but
Soares considered that the shelled, toasted product had a better taste.
Portuguese women in Brazil made several confections from mixtures of
sugar and groundnut.
Post-Columbian historical narratives 29
Following the conquest of the Aztecs, the conquistadors sent to the
emperor many reports of the natural resources in Mexico. These docu-
ments, in the Spanish archives, are generally unavailable to researchers, so
that access to information concerning possible distribution and use of
groundnut in Aztec agriculture is limited. Friar B. de Sahagun (1820-30),
compiler of an encyclopedia in the Nahuatl language (1558-1566), men-
tioned a folk-medicine use of tlalcacautl (Nahuatl for groundnut) but did
not list it among the principal food plants of central Mexico. Nor is it
among records of the tributes Montezuma extracted from tribes conquered
by the Aztecs. The compound name tlalcacautl is cited as evidence of its
late arrival in Mexico. Krapovickas (1968) suggested that the introduction
was probably a subspecies hypogaea type grown in the Antilles. Despite
recent archaeological evidence for the relative antiquity of cultivation, the
absence of other Arachis species in Mexico is substantive evidence that the
groundnut is not native to the area, nor was it domesticated there.
2.3.2 Selected seventeenth century narratives

Early in the seventeenth century, descriptions and illustrations of ground-
nut appeared regularly in the European literature and botanical gardens
grew the plant as a curiosity. Many natural historians were annotators,
compilers, copiers, editors, illustrators and translators who systematized
the observations of others but rarely saw the plants (or animals) whose
descriptions and figures they placed in their folios. In contrast are the field
studies of four naturalists: Marcgrave de Liebstad (1648, 1658), Cobo
(1653), Tertre (1654) and Labat (1697, 1742), whose descriptions and
figures came from material they observed and collected in natural habitats.
An English translation of Monardes' book appeared in 1577 and editions in
several other languages quickly followed. The first figure of the beaked
pods of a Brazilian cuItigen with two or three seed cavities is that of Jan de
Laet, naturalist, editor and managing director of the Dutch West Indies
Company. The company's ship captains brought him plant collections from
many parts of the New World. However, Laet's description (1625, 1640)
follows Lery's text (1578).
After the Dutch wrested control of the northeastern Brazilian coast from
the Spanish Empire in 1630, Governor-General J. Maurice von Nassau-
Siegen commissioned a scientific expedition (1638-1641) of Pernambuco
by his personal physician WillemPiso and his friend George Marcgrave of
Liebstad. A naturalist, astronomer and geographer, Marcgrave collected
the plants reported in their books. After Marcgrave's death in 1640,
Jan de Laet became his literary executor and edited and published the
Marcgrave-Piso notes and figures. Their Natural History of Brazil includes
Marcgrave's eight books (1648, 1658). Marcgrave described the groundnut
(mandubi) plant in some detail, illustrating the two-seeded fruits, quadrifo-
liate leaves with opposite leaflets, and flowers in the axillary position, but
he erroneously placed the fruits growing on the roots. In a second edition
(1658) issued by Piso, Marcgrave's description reappears but two pods of a
different botanical variety (apparently copied from an edition of Laet's
1625 book) were added to the illustration. Cobo (1653), who also thought
the plant produced fruit on the roots as in sweet potato, gave essentially
the same description.
The first French botanist in the Antilles, the priest J.B. du Tertre,
described the groundnut plant brought to him by the Carib Indians in
Guadeloupe (Tertre, 1654). In the location, configuration and morphology
of organs, however, Tertre's figure bears a striking likeness to that of
Marcgrave (1648), and Tertre's three-segmented pod is that figured in
Marcgrave's revised work (1658).
Labat, another priest, who lived in the French Antilles for 12 years
(1693-1705), gave a remarkable description of groundnut in his New
Voyage to the Isles of America, written in 1697:
the fruits came from a plant hardly a foot tall, a creeper with many
slender stems ... leaves resembling sweet clover, and nasturtium-
coloured flowers .... The short life of the flowers is due to their
shrivelling in the sun. The fruit is found in the earth ... attached by
filaments to hairs that the roots put out [sic] from stems distributed on
the surface of the earth, where they enter and produce pods.
Pods were 2.7-4.0 cm in length and 0.9-1.35 cm in diameter. Labat (1697,
1742) described shell reticulation, seed shape and number per pod, and the
testae, and he recorded different culinary uses of the seed. He found some
indigestibility from eating the seed raw and claimed that 'roasted ground-
nut stimulates the appetite and thirst'.
In view of Labat's comment concerning groundnut's persistence of
volunteering in fields after harvesting, the type he saw may have possessed
appreciable fresh-seed dormancy. Labat's figure (1697, 1742) reverses the
illustration in du Tertre (1654) and both appear identi,cal with the branch
and opened pod figured in Marcgrave (1658).
British naturalists came late to the West Indies. Sir Hans Sloane visited
Jamaica, Barbados and St. Kitts in 1687-88. Sloane's catalogue (1696)
describes groundnut and, more importantly for this review, he cited at least
14 authors who recorded groundnut in or from the Americas (Hammons,
1982). His frequent references to the plant show the prevalence and wide
use of groundnut as a food crop in the islands he visited.
By the beginning of the eighteenth century, at least 20 botanical or
historical works describing the groundnut were in print and in wide circu-
lation among historians. Besides the universal Latin editions, translations
appeared in most of the major languages of western Europe. The main
exception is the important work of Soares de Souza, written in 1587 but not
published until 1825. All these authors knew that groundnut was native to
the Americas.
Origin of groundnut 31
2.4 INDIAN VERNACULAR NAMES
The distribution and variety of local names of the groundnut are important
in the study of its origin. The Spanish introduced the name mani from the
West Indies and substituted it for other Indian names in Spanish America,
except Mexico. There they accepted the alternative Nahuatl word caca-
huate, modified it and took it to Spain as cacahuete. The Portuguese
amendoim stems from many cognate names still used in Brazil in the late
twentieth century.
Cobo (1653) listed Indian vernacular names to document groundnut's
diffusion in pre-discovery America: 'This root [sic] is called mani in the
language of Hispaniola, Mexicans call it cacahuate, and the Peruvian
Indians call it inchic in the Quichua language and chocapa in Aymara.'
Marcgrave (1648) used the Brazilian Indian mandubi, and, citing
Monardes (1574), 'anchic of Peru, the same is called manf in Spanish.'
Tertre (1654) coined the French name pistache because of the seed shape
and its taste similar to the European pistachio. Sloane (1696) compiled
eleven common name synonyms: manobf, mandovy, munduvi, anchic,
ibimanf, manf, ynchic, pistache, mandues, earthnuts or pindalls.
Krapovickas (1968) successfully associated the latin names of many
earlier authors with the subspecific variation within Arachis hypogaea and
then associated the ethnic, linguistic and geographic centres of origin of the
cultivated groundnut. He also (1968, 1973) correlated Amerindian names
with groundnut diffusion throughout South America. In the Tupi-Guarani
region, he found numerous variants of manduvi. Vernacular names
showed little linguistic affinity in the remainder of the continent. The
greatest diversity of Indian names occurs on the eastern slopes of the
Andes, where Bolivia and Peru join and where manduvi variants are
interspersed with other names of Arawak affinity.
The Arawaks inhabited a vast area extending from the Caribbean to the
heart of South America as far as the Bolivian border with the Chaco
(Hammons, 1982). The presumed centre of origin of A. hypogaea is the
region where Arawak linguistic influences predominate and Krapovickas
(1968) presented a tenable hypothesis of Arawak responsibility for the
spread of groundnut from such a centre to the Caribbean basin.
2.S ORIGIN OF GROUNDNUT
2.S.1 Geographical origin of genus Arachis

The genus Arachis L. comprises a large and diverse group of diploid and
tetraploid taxa native only to South America. There Arachis ranges geo-
graphically from the equator near the mouth of the Amazon to 34° S
latitude on the northern bank of the Rio de la Plata in Uruguay, and
extends westward from the Atlantic to the Parana and the eastern foothills
of the Andes (35-66° W longitude). The northern boundary is usually
marked by the southern extent of the Amazonian rainforest, except where
Arachis spp. may occur where the forest is penetrated by the more open
vegetation of the cerra do breaks. Gregory et al. (1980) theorize that the
inferred centre of distribution for the genus is the Brazilian Planalto Ellipse
and that the only plausible natural agent of distribution is the downward
flow of soil and water. Biological dispersal mechanisms are very poor, with
the annual dispersion rate limited to a radius varying from several centi-
metres to a few metres.
Both Krapovickas (1968, 1973) and Gregory et al. (1980) postulated a
Plan alto profile from Corumba to Joazeiro, Brazil, as the centre from
which distribution of Arachis arose. Half of the eight sections into which
the genus naturally divides botanically occur there.
Although 50-70 species may exist in the genus, only 23 species, one
variety and one hybrid are validly published (Stalker, 1990). All are
indigenous to the area east of the Andes and south of the Amazon
rainforest.
2.5.2 Origin of Arachis hypogaea

Linnaeus (1753) described the first Arachis in his Species Plantarium. Five
species, each from different sections, are now cultivated. A. hypogaea L.,
the principal cultigen, and A. villosulicarpa Hoehne are grown for their
edible seeds and were improved by the indigenous peoples of South
America. Recently, forms of A. glabrata Benth., A. repens Handro, and
A. pintoi Krap. et Greg. nom. nud. were purposely adapted for grazing use
and released. A. rep ens is also in use as the principal ground cover (lawn)
at the Museum of Natural History in Sao Paulo and at the Clube Praia das
Cigarras resort, 7 km west of San Sebastiao, Brazil (R. Hammons, unpub-
lished field party notes, 2 June 1968). The low genetic affinity of these five
species shows their independent domestication. This is especially interest-
ing for A. hypogaea and A. villosulicarpa, both cultivated for their qualities
for human nutrition: A. hypogaea is widely spread throughout the world
but A. villosulicarpa has limited use only by the Indians of the Rondonia
area of the Mato Grosso, Brazil (Krapovickas, 1968). (Conversely, one
may argue that A. hypogaea is the only domesticant, whereas the others
are cultivated, i.e. they can survive in the wild.) Arachis hypogaea
(2n = 4x = 40) is a member of section Arachis nom. nud., along with
its tetraploid progenitor, A. monticola Krap. et Rig. (Stalker, 1990).
Identification of the diploid ancestors is inconclusive. Krapovickas (1968)
proposed southern Bolivia and northwestern Argentina for the origin of A.
hypogaea. The area is an important centre of diversity for subsp. hypogaea.
Although A. hypogaea is a largely self-pollinating, annual, herbaceous
legume, earlier work of Krapovickas and Rigoni (1960) showed that the
Dispersal 33
great variability of groundnut in the Guarani region (the basins of
the Parana-Paraguay river systems) resulted from a hybrid swarm of the
original prototypes with natural crossing occurring freely. An extension of
this hypothesis, with different prototypes, to the rest of South America
gave rise, according to Krapovickas (1968), to the patterns of variability
that he and Gregory et al. (1973, 1980) delineated as distinct gene centres.
The eastern foothills of the Andes constitute a wide range of ecologically
distinct environments. In this proposed centre of origin, where small-scale
cultivation is practised, Krapovickas (1968) cited the diversity of uses as
further evidence of the antiquity of groundnut. Immature to fully ripe
seeds are eaten raw or cooked. They may be boiled, broiled, roasted,
toasted, fried or ground into a paste for mixing with other food. Soups may
occasionally contain boiled whole young pods. The common refreshing
non-alcoholic drink, chica de manf, is popular there and soap is made from
oil pressed from groundnut seed. (Note: in western Brazil, chica is an
alcoholic drink.)
A Bolivian origin is also supported by the wide range in seed and pod
morphology documented there, for example, by Cardenas (1969).
However, no one is certain of the exact origin of groundnut. All the
linguistic, cytological, genetic, morphological, biochemical and geographic
data indicate that the eastern foothills of the Andes is the region where A.
hypogaea originated. Information from the large reservoir of groundnut
landraces collected over a vast area of South America, beginning in 1959,
should provide new insight on genetic homologies, biochemical affinities,
original habitat, domestication and dispersal.
2.6 DISPERSAL
Prior to the early 1500s, groundnut was unknown outside the Americas.
Worldwide distribution of at least two distinct forms - a two-seeded
Brazilian and a three-seeded Peruvian - took place comparatively soon
after the discovery of the New World (Dubard, 1906). Many authorities
credit the Portuguese with enriching African agriculture by introducing
groundnut there from Brazil, carrying it subsequently to the Malabar coast
of south-western India, and possibly to other lands. However, I found no
documentation that they did intentionally introduce the seed to those
lands.
Gibbons et al. (1972) observed that cultivar clusters in Africa of subsp.
Jastigiata var. vulgaris represented both the Guarani region and the region
of the eastern slopes of the Andes in Bolivia and parts of western Brazil
(Gregory et al. 1973). As the latter authors note:
It is also fairly plain that the peanuts of the eastern slopes of the
Andes must have reached Africa from Portuguese boats plying the
Amazon and not from the immediate interior back of the northeast
coast of Brazil.
The Peruvian type A. hypogaea var. hirsuta went to the western Pacific,
to China, Indonesia (Java), and to Madagascar. Dubard (1906) docu-
mented the concurrence in morphology and configuration of random pod
samples from the latter three places and between these and the 'hump-
backed' material found in tombs at Anc6n, Peru. Their most plausible path
was up the west coast from Peru to Mexico, thence across the Pacific as an
item of trade on ship crossings that were regularly scheduled between
Acapulco and Manilla (the Philippines) for the 250 years prior to 1815
(Krapovickas, 1968). .
In the African and Asian lands where it was carried, the groundnut
readapted for environmental and specialized agricultural requirements.
Precisely how and when groundnut entered the United States is unknown.
Indeed, there may be instances of its being introduced indirectly from the
European farming tradition into colonial America. Nor can one rule out
introduction directly from its native South America and secondary
Mesoamerican agriculture. Any such technological borrowing could have
been mediated by contacts between landowners, the inspection of fields
and botanical gardens, or the reports of travellers. Probable locations for
the transfer of seed and husbandry information could be the Iberian
peninsula, southern France or Italy. The diffusion of technology from the
Old World back to the New may always remain partly conjecture. The
anonymity of the event should not devalue the achievement.
Burkhill (1901) cited Clusius (1605) as saying that:
slavers took as food for their captives on the voyage from the Guinea
Coast of Lisbon, roots of the sweet potato, 'besides certain nuts',
which Sloane identifies as fruits of Arachis.
But, as Burkhill stressed, Clusius' observation places Sloane's identifi-
cation in question (Hammons, 1982).
A small-podded genotype with a spreading habit of growth was the
earliest form successfully introduced into commerce in the south-eastern
United States. It is a long-season groundnut, possibly from Africa (Hig-
gins, 1951), and its pod and seed morphology, aspect and branching
pattern agree with phenotypes described and illustrated in the West Indies
by Tertre (1654) and Labat (1697, 1742). Thus, direct introduction from
the Caribbean cannot be excluded (Hammons, 1982).
The Guarani region of north-eastern Argentina, Paraguay and south-
western Brazil is the centre of variation for the spanish type (subsp.
Jastigiata var. vulgaris) and it was distributed from this region. According
to Krapovickas (1968), F.L. Gilli and G. Xuarez documented its introduc-
tion into Europe: seed from Brazil came in 1784 to Don Jose Campos in
Lisbon, who sent a portion on to Rome. Tabares de Ulloa (1799) spread
Ethnography 35
the type in Valencia. From there, Lucien Bonaparte carried it to the south
of France in 1801 (Burkhill, 1901; Andre, 1932). The Spanish, who culti-
vated the crop both for cooking oil and for use in preparing a chocolate-
covered confection, extracted the first oil in Europe (Dubec, 1822).
The spanish type is a small two-seeded form adapted to adverse environ-
mental conditions; it has a shorter growing cycle and lacks appreciable
fresh-seed dormancy. T.B. Rowland successfully introduced it from
Malaga, Spain, in 1871 to Norfolk, Virginia (USA), where he distributed
seed without cost among planters (Anon., 1918b).
Gregory et at. (1980) associated the virginia type (subsp. hypogaea var.
hypogaea) with both the Bolivian and Amazonian geographical regions,
but its origin needs further clarification. Extensive secondary variations
occurred in Africa (Gibbons et at. 1972). McClenny (1935) has placed
its cultivation in Virginia as early as 1844; other records suggest that its
introduction followed the pioneer work of Rowland. The jumbo virginia of
United States commerce may be from a chance hybrid (Anon., 1918a).
Recently, Williams (1991 b) presented ethnobotanical evidence for the
Bolivian origin of the valencia type (subsp. fastigiata var. fastigiata) , which
Krapovickas (1968) postulated spread throughout the world from Paraguay
and central Brazil. Williams (1991a, b) investigated the farming practices
of indigenous growers, cultivating six distinct landraces along the Rio Beni
using age-old agricultural procedures. Groundnut is cropped on exposed
riverine sandbars during the low-water season. A selected portion of the
harvest is replanted in upland gardens as a seed crop. The cropping system
exerts strong selection pressure for the sub specific valencia characters of
earliness, sequential flowering, loss of seed dormancy, and strong pegs.
Bees are frequent visitors to the flowers. Such data, with the archaeological
and historical evidence, and the diversity of valencia landraces, further
support the notion that ancient people living in this region also developed
the valencia botanical type (Williams, 1991a, b).
Although Dubard (1906) described the fruit, Beattie (1911) apparently
chose the name 'valencia' to designate an introduction into the United
States from Valencia, Spain. Soon 'valencia' became a generic name for
biotypes with similar pod configuration and a unique arrangement of
vegetative and reproductive branches.
'Waspada', with approximately 50% earlier maturity, was the first culti-
var introduced to achieve a specific agricultural goal: a shorter growing
season. Brought to Java in 1875, the cultivar eventually replaced the
common form that matured in eight to nine months (Holle, 1877).
2.7 ETHNOGRAPHY
Ethnographic investigations among the major Indian tribes of South

America document the widespread cultivation and use of groundnut as a
food crop throughout much of the continent. The studies also provide
indirect evidence supporting its domestication long before the Spanish
conquest. In the seven volumes of the Handbook of South American
Indians, Steward and his collaborators (1943-1959) traced groundnut dis-
persion through records of food plants sown and harvested by native
peoples. The following information summarizes the major findings.
The Spaniards found more than 40 new food plants - including ground-
nut - for pre-Columbian civilization in the fertile highland basins and
coastal valleys of the central Andes. Metraux (1942) reported that the
Spaniards who penetrated eastern Bolivia with Gonzalo de Solis Holguin,
Governor of Santa Cruz de la Sierra, in 1617 and again in 1624 were
amazed at the large plantations where the Mojo (c. 15° Sand 65° W) and
Baure tribes grew groundnut, preferably along sandy river beaches of the
Mamore's tributaries in the upper Amazon river system. Some 375 years
later, Williams (1991a, b) provided the first detailed ethnobotanical exam-
ination of this agricultural practice, in areas to the west in a diverse
rainforest environment on the Rio Beni.
Sixteenth century Indians in Paraguay grew groundnut as a main crop for
trade in the markets of Asuncion. Throughout the vast tropical forest many
indigenous aborigine tribes grew groundnut as a staple crop on farms in
natural or manmade clearings. In central Brazil, large gardens guaranteed
an economy of abundance for the Tapirape tribe (c. 10° S, 52° W). Here, as
Soares (1587) observed for the east coastal area, women planted and
harvested the crop. In the Caribbean basin, the Arawak tribe grew ground-
nut with the aid of irrigation (Stewart, 1943-1959). Irrigation was always
practised where groundnut was grown in the arid climate of coastal Peru
(Cohen, 1977; Burger, 1989).
2.8 EARLY INDUSTRIAL DEVELOPMENTS
Tabares de Ulloa (1799), later bishop of Valencia, invented the first

machine to shell groundnut. A description of this device appeared in an
1805 supplemental issue of Rozier's Traite General d'Agriculture.
2.8.1 Beginning of the groundnut oil industry

Worldwide shortage of oil in Europe during early decades of the nine-
teenth century led to industrial development of the groundnut oil industry.
Conflicting claims, that fail to differentiate between samples imported for
experimental trial and shipments to mills for crushing, cloud the question
of credit for initiating the commercialization of groundnut production in
Africa for export, and when this occurred. Once trade began, however,
exports increased at a rapid rate. For example, Gambian exports to Britain
Early industrial developments 37
went from 213 baskets in 1834 to 47 tons in 1835, and jumped to thousands
of tons by the early 1840s (Brooks, 1975).
America recorded its first imports from Gambia in 1835 and American
purchases dominated the Gambian market from 1837 to 1841. The
American interest in the West African groundnut differed from that of the
Europeans: the latter desired the oil product, wheras Americans relished
the roasted groundnut (Brooks, 1975).
French industrialists also entered the ground nut trade during the 1830s.
Fliickiger and Hanbury (1879) credit Jaubert, a French colonist and trader
at Goree near Cape Verde, with first suggesting commercialization by
sending an oil sample to Marseilles, France, in 1833. He initiated the
industry with a shipment of 722 kg to Marseilles in 1840, when France
reduced the tariff on groundnut (Brooks, 1975). Following the import of a
large tonnage from Cape Verde in 1848, groundnut became a most import-
ant raw material for the French oil industry for more than a century
(Schlossstein, 1918). In the first decade of the twentieth century, France
led the world in tonnage of groundnut crushed for oil, followed in volume
by Germany, Holland, Austria and then England (Schlossstein, 1918).
In the United States, initial expansions in land area cropped and in
production came in response to the increased need for oil for various uses
in times of shortages due to war and other causes. Following the Civil War,
the crop area doubled and trebled between 1865 and 1870. Other spurts
coincided with the Spanish-American War and World Wars I and II.
Moreover, as the boll weevil destroyed the cotton crop in the southern
states early in this century, farmers turned to ground nut for economic
relief.
The first groundnut pod polisher, invented by T.B. Rowland in the
1870s, was operated by draught animal (mule) power. It embodied mech-
anical principles used for all subsequent polishers (Anon., 1918b).
2.8.2 Peanut butter

Until the late 1890s consumption of groundnut in America was mostly
roasted-in-the-shell. However, in 1894, Dr. John Harvey Kellogg, MD, an
eminent health reform advocate and director of the Sanitarium [sic] in
Battle Creek, Michigan, developed and introduced peanut butter as a
nutritious, easily digested health food for patients (Powell, 1956). Kellogg,
who observed a strictly vegetarian diet, obtained the first two patents for
making peanut butter (US Pat.No. 580787, issued 13 April 1897; US
Pat.No. 604493, issued 24 May 1898). To promote his health food ideas,
Kellogg elected not to enforce his patent rights and several brands of
peanut butter were on the market by 1899.
A former employee of the Sanitarium, Joseph Lambert, began to manu-
facture and market three machines - a roaster, a blancher and a hand nut-
grinding mill - for making peanut butter in 1896 (Grohens, 1920).
2.8.3 Subsequent development
The commercialization of groundnut food products expanded rapidly
thereafter. Scientists developed new cultivars and cultural practices, and
improved agricultural and industrial machinery became available for all
phases of groundnut production, processing and manufacturing.
Periodically, groundnut oil is proposed for use as a substitute or extender
for diesel fuel. When Rudolf Diesel demonstrated the engine that bears his
name at the Paris exposition of 1890, it was powered entirely with groundnut
oil (Nitske and Wilson, 1965). This practice has not been feasible sub-
sequently, both economically and mechanically (Hammons, 1981).
Without the primitive plant selectors who fashioned the groundnut into
useful sources of oil and protein, we would not have this crop whose fruit can
be eaten raw or cooked and whose seed can be stored for future use and for
seed. Who were these early selectors? Anthropologists and ethnographers
say that they probably were women - women who gathered the seed and
tended the crop, as Soares de Souza (1587) and Williams (1991a) have
noted. As Burton (1981) comments, they lacked a knowledge of genetics
and reproductive behaviour in plants, of mutations and metabolic pathways,
of pathogens or pests and their control; they had neither written language,
libraries nor computers. But they had their plants, though undoubtedly a
much more restricted germplasm base than groundnut breeders would wish,
as Williams (1991a, b) among others, has illustrated. They knew their plant
material from living with it: they were motivated because their lives
depended on their own success. And they had time on their hands.
What did they do? They used a rudimentary but primary activity of all
plant breeding procedures - selection - to choose the material best adapted
for their goal: food for survival. They increased the yield, perhaps many
times, from the basic stocks inherited from their predecessors. As the crop
dispersed over the millennia from the centres of origin, new selectors
sought or preserved changes - for environmental adaptation, for resistance
to pests and diseases, and for more nutritious qualities - essential for
genetic diversity under continued domestication.
Thus, by the time of the European encounter with the Americas,
Arachis hypogaea already existed in many landraces, distributed and under
cultivation throughout much of South and Central America, and the
Antilles and other islands of the Caribbean basin. The full extent of its
dispersion may soon became known as scholars see the narrative reports
kept in the royal Spanish archives. Recent breakthroughs in decoding the
complex of hieroglyphs forming the written system of the Mayas may
reveal a broader use of groundnut in agriculture and the diet in Mexico
than that recorded by Sahagun in the Nahuatl language in 1558-1566, but
suppressed until 1820.
Additional archaeological evidence could come as Alva (1988) com-
pletes the inventory of Moche culture material from the tomb near Sipi'm,
References 39
Peru, and further Moche or Classic Mayan records will likely be revealed
by the renewed interest in the prehistory of native peoples in
Mesoamerica.
The groundnut, by the very nature of its underground fruit placement,
was not among the colourful New World crops that excited the sixteenth
century European chef; moreover, except in Spain, it was ill-adapted to
Europe's agricultural requirements. Although the plant soon became a
curiosity in Europe's botanical gardens, it was poorly understood long after
Linnaeus (1753) formalized its name, as Gregory et al. (1973) have well
described.
The food historian has yet to document fully the importance and diver-
sity of groundnut in the diet of native peoples prior to their encounter with
Europeans. The groundnut was overshadowed by other novel foods:
maize, potatoes, green beans, pineapple and tomato. Even the capsicum
peppers made their way more quickly into world cuisine than the nutritious
groundnut.
The Quincentennial in 1992 of the encounter between the two worlds
may yet focus renewed attention on the groundnut - one of the 13 crops
that stand between humans and starvation.
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Oviedo y Valdes, G.F. de (1851) Cronica de las Indias, Salamanca, lib. 23, cap. 12, p. 193.
(Completed work issuedby the Spanish Royal Academy of History in 1851.)
Powell, H.B. (1956) The Original has this Signature - W.K. Kellogg, Prentice-Hall, Inc.,
Englewood Cliffs, New Jersey.
Sahagun, B. de (1820-30) Historia General de las cosas de Neuva Espana. Republished 1956,
A.M. Garibay-K., Editorial Porrua, S.A., Mexico, D.F., 4 vols; v. 3, tlalcacuatl, p. 173.
Original Ms in Nahuatl, 1558-1566.
Schlossstein, H. (1918) The peanut situation as affected by world events. The Peanut
Promoter, 1, (4), 32-33.
Schmidt, U. (Faber, U.) (1567) Weltbuchs von Newen erfundnen Landtschafften: Warhafftige
... , Original German edn 2 vols., Frankfort am Main. Translation (1891) from the
original German text: The Conquest of the River Plate (1535-1555), Part I. Voyage of
Ulrich Schmidt to the Rivers La Plata and Paraguai, (cd. L.L. Dominguez), Hakluyt
Society Publication, vol. 81, London, pp. 25, 37, 40-41, 63.
Sloane, H. (1696) Catalogus Plantarium que in Insula Jamaica, etc., D. Brown, London, pp.
72-73,221.
Smith, C.E., Jr (1967) Plant remains, in The Prehistory of the Tehaucan Valley (ed. D.S.
Byers), University of Texas, Austin, Texas, vol. 1, pp. 220-225.
Soares de Souza, G. (1587, 1825, 1851) Tratado Descriptivo do Brasil em J587, first published
1825, in Colle«ao de Noticias para a Historia c Geographia das Nac6es Ultramarinas, que
vivem nos Dominios Portuguezes, Academie de Sciencias, Lisbon. Republished 1851,
separate issue, Revista trimonsal do Instituto Historico e Geographico do Brasil, 14, 1-423.
Amendoim, cap. 47, pp. 173-176.
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Macmillan, New York and London.
Stalker, H.T. (1990) A morphological appraisal of wild species in scction Arachis of peanuts.
Peanut Science, 17, 117-122.
Stewart, J.H. (ed.) (1943-1959) Handbook of South American Indians, Bulletin 143, Bureau
of American Ethnology, 7 vols, Smithsonian Institution, Washington.
Tabares de Ulloa, F. (1799) del Cacahuate 6 Manf de America, Semananario de Agricultura y
Artes, 5 (123), 289-294.
42 The origin and history of the ground nut
Tertre, J.B. du (1654) Histoire Gentirale des Isles . .. et autres dans ['Amerique, Paris.
(1667-71) rev. edn, 4 vols, Histoire Generale des Antilies Habirees les Frant;ois, T. Jolly,
Paris, v. 2, p. 121 and plate.
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Williams, D.E. (1991a) Peanuts and peanut farmers of the Rio Beni: Traditional crop genetic
resource management in the Bolivian Amazon, PhD dissertation, City University of New
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(Abstract), American Peanut Research and Education Society, Proceedings, 23, p. 22.
CHAPTER 3
Botany - morphology and anatomy

V. Ramanatha Rao and U.R. Murty
3.1 INTRODUCTION
Groundnut (Arachis hypogaea L.) is one of the world's major food legume
crops. It originated in South America, where the genus Arachis is widely
distributed. The natural distribution of all the Arachis species is confined to
Argentina, Bolivia, Brazil, Paraguay and Uruguay (Krapovickas, 1973;
Krapovickas and Rigoni, 1957; Ramanatha Rao, 1987). A. hypogaea is not
known to occur in the wild state. Groundnut is presently cultivated in over
80 countries from 40° N to 40° S in tropical and warm temperate regions of
the world. The origin and evolution of the genus as well as of the cultivated
species are fairly well known but more evidence is required to make
explicit conclusions. Though considerable information on morphology,
cytology and genetics has been accumulated along with information on the
distribution of the species in the genus, the taxonomic treatment is still
incomplete. One of the major reasons for this is the discovery of additional
'species' after every collecting mission to centres of diversity of the genus
(Valls et al., 1985). The morphology, development and anatomy of Arachis
presented problems to groundnut workers.
This chapter attempts to review these topics and to clarify certain
ambiguities. However, most of the discussion is on the cultivated ground-
nut rather than on its wild relatives. Information on origin, evolution and
taxonomy is limited to the extent that it is relative to morphology and
development.
3.2 DESCRIPTION OF THE GENUS
Arachis is a perennial or annual legume with three or four leaflets, stipulate

leaves, papilionate flowers, tubular hypanthium and subterranean fruits.
The 'peg', which is an expanded intercalary meristem at the base of the
1994 by Chapman & Hall. London. ISBN 0412408201.
44 Botany - morphology and anatomy
basal ovule (Gregory et at., 1973), is unique to the genus. The expansion of
the intercalary meristem results in a lomentiform carpel of one to five
segments, each containing a single seed with two very large cotyledons and
a straight embryo. A general description of Arachis is as follows (after
Bentham and Hooker, 1862):
Seeds with thick and fleshy cotyledons; short radicle growing into a
well-developed taproot; plants low suberect herbs, often prostrate
and even creeping; leaves abruptly bipinnate, tetrafoliolate leaves
with two pairs of opposite leaflets, rarely trifoliolate, exstipellate,
stipules adnate to the petiole at the base. Flowers, crowded in simple
or compound monopodium, look like dense axillary spikes, sessile in
the leaf axil, or very shortly pedicellate. Calyx lobes five, often
dentate, calyx tube filiform, lobes membranaceous, the four upper
ones connate, the lower one slender, separate. Petals and stamens
inserted at the apex of the tube, standard oblong, obovate to suborbi-
cular, wings oblong, free, keel incurved, prostrate. All stamens
connate to form a closed staminal tube, stamens 10, usually one
absent, anthers alternate, elongate, subbasifixed, the alternate ones
versatile. Ovary subsessile towards the base of the calyx tube, usually
aerial, occasionally subterranean, two- to three-ovuled. When the
flower withers and falls away, the ovary shows a stalk, which
elongates and becomes reflexed and rigid and the ovary is continuous
with the same, acute at the apex; style long, filiform with a minute
terminal stigma; the pod ripens inside the soil, oblong, thick, indehis-
cent, subtorulose, articulate or non-articulate. Seeds one to three,
irregularly ovoid, rich in oil and protein.
3.3 DESCRIPTION OF THE SPECIES
In this section, brief morphological descriptions of the valid Arachis wild

species are given. The authors feel that these descriptions serve as useful
guides to Arachis species because a taxonomic monograph of the genus is
still lacking. Much of what is given below is an English translation of the
original descriptions given by the respective authors (the protologues)
and is adapted from Ramanatha Rao (1988). Where appropriate, some
additional information is also provided. Most of the measurements given
are averages.
Besides the valid names, a few nomina nuda or nomina inedita are
used in the literature. Some of these are A. chacoense (= A. chacoensis) ,
A. correntina, A. cardenasii, A. pintoi, A. stenosperma (= A. stenocarpa),
A. sytvestris, A. ipaensis, A. spegazzini, A. oteroi, A. lignosa, A. duranen-
sis and A. macedoi (Ramanatha Rao, 1988; Resslar, 1980). These are
taxonomically invalid and should not be used; if used they should accomp-
any the collector numbers. The 22 species described so far are listed in
Description of the species 45
TABLE 3.1 Valid Arachis epithets with citation and botanical assignment
Species Author citation Section/

Series
A. batizocoi Krap. et Greg. Krapovickas et al., 1974 Al

A. villosa Benth. Bentham, 1841 A2
A. diogoi Hoehne Hoehne, 1919 A2
A. helodes Mart. ex Krap. et Greg. Krapovickas and Rigoni, 1957 A2
A. hypogaea L. Linnaeus, 1753 A3
A. nambyquarae Hoehne Hoehne, 1922 A3
A. monticola Krap. et Rig. Krapovickas and Rigoni, 1957 A3
A. tuberosa Benth. Bentham, 1841 El
A. guaranitica Chod. et HassI. Chodat and Hassler, 1904 El
A. paraguariensis Chod. et HassI. Chodat and Hassler, 1904 E2
A. benthamii Handro Handro, 1958 E2
A. martii Handro Handro, 1958 E2
A. rigonii Krap. et Greg. Krapovickas and Gregory, 1960 E3
A. repens Handro Handro, 1958 C
A. burkartii Handro Handro, 1958 Rl
A. glabrata Benth. Bentham, 1841 R2
A. hagenbeckii Harms Kuntze, 1898 R2
A. prostrata Benth. Bentham, 1841 EX
A. marginata Gard. Gardner, 1842 EX
A. villosulicarpa Hoehne Hoehne, 1944 EX
A. lutescens Krap. et Rig. Krapovickas and Rigoni, 1957 EX
A. pusilla Benth. Bentham, 1841 T
Table 3.1. Though some workers include A. angustifolia (Chad. et HassI.)

Killip (in Hoehne, 1940) as one of the described species (Smartt, 1990),
the validity of the name as given in the literature is questionable.
A. X batizogaea Krap. et Fern., an experimental hybrid origin from a cross
between A. hypogaea and A. batizocoi (Krapovickas et al., 1974) is also
not included. A. correntina (Burk) Krap. et Greg. nom. nud. is A. villosa
var. correntina Burkart, but needs validation according to the International
Code of Botanical Nomenclature. It has been suggested that A. namby-
quarae should be regarded as a variety of A. hypogaea (John et al., 1954)
but its description is not given here because it is simply a form of
A. hypogaea (Smartt and Stalker, 1982) and not a distinct species.
Arachis batizocoi Krap. et Greg.

Annual, prostrate herb; erect main stem, up to 15 cm, hairy; tetrafoliolate;
outer surface of stipules hairy, 2.5 cm x 1 cm, mostly adnate to the petiole;
petiole 5 cm long on main stem, 4 cm on branches, hairy; leaflets elliptic on
main stem, 3 cm x 2 cm, orbicular on branches, 2.5 cm in diameter, hairy
underneath, ciliate margins, tip acute on main stem, rounded, obtuse or
mucronate on branches, coriaceous to brittle; corolla yellow, generally
with pink or purple blush on the back of the standard; hypanthium long;
pods articulate, 10 mm x 5 mm, with prominent beak, seeds light tan,
Smm x 4mm.
Arachis villosa Benth.

Perennial, prostrate; stems sometimes angular, villous, 5-14 cm tall; little
branching, primaries 40-100 cm or longer; tetrafoliolate; stipules elongate,
falcate or lanceolate, 1-2 cm long, rigid, very hairy, shortly adnate;
petioles 2-3 cm, hairy; rachis 3-7 mm; leaflets broadly ovate, rigid, mucro-
nate, emarginate, short dense hairs above, sparsely hairy below, O.S-
1.5 cm x 0.5-1.0 cm, generally apical leaflets longer than basal; thread-like
hypanthium, 2-4 cm, hairy; corolla glabrous; calyx upper lip tri- or bila-
biate, 6-7 mm, lower lip wide, entire or toothed; standard yellow or
yellow-orange, 12-15 mm in diameter; ovary with two ovules; pegs up to
20 cm, two or three articles of pod separated by isthmus which rots after
seeds mature, isthmus 2 cm, articles 11-16 mm x 7-10 mm, slightly ridged,
beak prominent; seeds reddish brown or tan, 10 mm x 5 mm.
Arachis diogoi Hoehne

Perennial; prostrate; stem angular, well-branched, dense interwoven white
hairs on stem, branches, leaves, stipules; internodes 4-6 cm long; tetrafo-
liolate; stipules adnate to the petiole up to 6 mm, long-acuminate, striate,
2 cm long; petioles 4-6 cm long; leaflets narrow-oblong to linear-oblong,
tapering to base and apex, base subrounded, apex acute, glabrous above,
thinly hairy below and on margins, 3-4 cm x 0.7-1.0 cm; flowers one to
four per axil, hypanthium 4-7 cm long, thin, hairy; calyx bilabiate, lobes
10 mm long, upper lip minutely three-toothed, lower lip entire, narrow-
acute; standard yellow or orange-yellow, suborbicular, 12 mm in diameter,
apex emarginate, base contracted in short claws, reflexed -above; wings
with rounded tip, subfalcate, dilatate above the middle, base clawed, eared
above the claw; keel narrow, base clawed, eared above, falcate, tip ros-
trate; stamens alternately short and long; anthers narrow up to 2.5 mm
long; pegs 3-S cm long, pericarp slightly reticulated, pods mostly uniarticu-
late, articles oblong, very slightly beaked.
Arachis helodes Mart. ex Krap. et Rig.

Perennial; taproot without rhizomes or stolons; stem highly branched,
prostrate; tetrafoliolate; stipules falcate, adnate to half of their length,
free part lanceolate-acuminate; petioles short; leaflets 0.5-1.1 cm x 0.4-
O.S cm, obovate-orbicular, emarginate, tip barely mucronate, margins
slightly thickened with few hairs, glabrous above and glabrous or subgla-
brous below; hypanthium thread-like, 3-6 cm long with scattered hairs;
calyx bilabiate, upper lip irregularly four-toothed, lower lip entire, linear-
acuminate, falcate, 7 mm long; standard lemon-yellow or yellow, orbicu-
lar, 10 mm in diameter; wing conspicuously eared; keel beaked, curved;
anthers eight, style filiform, longer than stamens, hairy at the tip, ovules
two; peg 5 cm long; articles 8-9 mm x 5-6 mm, pericarp thin, fragile,
hardly reticulate.
Arachis monticola Krap. et Rig.

Annual; taproot with profuse lateral roots; stem erect, radially symmetri-
cal, 30 cm tall, lateral branches distichous, creeping, hairy; stipules sub-
falcate, adnate to the petiole for one third of their length; leaflets
oblong-obovate; flowers up to five per axil; hypanthium thread-like, hairy,
5 cm long; calyx bilabiate, upper lip irregularly four-toothed, lower lip
entire, 10 mm long; standard yellow, suborbicular, 15 mm x 17 mm, keel
falcate, 12 mm long; anthers eight of which four are oblong, basifixed and
four small, globose and dorsifixed, style filiform, top hairy; ovules two;
pegs 10 cm long; pods biarticulate, articles 13 mm x 8 mm, isthmus 2-7 cm
long, pericarp reticulate, beaked; seeds light tan, 10 mm x 5 mm.
A. monticola is regarded as subspecies of A. hypogaea, because of the
high level of cross-compatibility and marked similarities between the two
(Smartt, 1964). It is very difficult to maintain this species in a pure form
due to the ease with which natural crossing occurs with A. hypogaea. The
fact that both alternate and sequential branching forms of A. monticola
occur (Gibbons, 1966) is a further confirmation of such a conclusion. A.
monticola could be regarded as either a subspecies or a botanical variety of
cultivated groundnut (Ramanatha Rao, 1988) but this needs to be
validated.
Arachis tuberosa Benth.

Perennial; roots with oblong woody tubers extending downward; stems
simple, very little branching, erect, rarely prostrate, softly adpressed,
villous, 20~30 cm high, angular, slightly sinuous; trifoliolate; stipules rigid,
striate, 2-3 cm long, adnate to the petiole from two thirds of their length to
almost to the tip, margins ciliate; petioles short; leaflets oblong, coriaceous
to rigid, marginate, reticulate, tip obtuse, base tapering, glabrous, 3.5-
4.0 cm X 1.0-1.3 cm; inflorescence subterranean, concentrated around the
crown of the stem or rarely at the base of branches; ovary underground but
hypanthium exposes flowers; hypanthium thread-like, erect, hairy, 5-7 cm
long; standard orange-yellow; pods mostly uniarticulate, 12 mm x 6 mm,
beak prominent, pericarp very fragile, slightly reticulated; seeds light tan,
8 mm X 5 mm, tend to lose viability when dry.
Arachis guaranitica Chod. et HassI.

Perennial; obconical root; erect stem, stems OrigInating from the root
tubers also erect; trifoliolate; closed leaf sheath forming tube covering
internodes; stipules long and congested, adnate to petiole, 3-6 cm long, tip
3-7 mm free; leaflets glabrous, linear-lanceolate, 5.0-9.5 x 1.5-3.0 cm,
marginal vein thick and midrib grooved, coriaceous, densely branched
striations; inflorescence almost entirely basal and arise in clusters around
the crown, flowers with subterranean ovary, flowers exposed by 6 cm long
hypanthium, hirsute; calyx limb bilabiate, upper lip four-toothed, hairy,
5 mm x 2.5 mm, lower lip entire, hairy and 7 mm wide; standard 18 mm
wide; wing emarginate; keel pointed, rostrate; stamens nine, five anthers
globose and four elongate; style thin, tip attenuate, hairy; pegs horizontal,
may grow up to 1 m at a depth of 5 cm; pods articulate, 10 mm x 5 mm,
with slight beak, seeds light tan, 8 mm x 4 mm, tend to lose viability when
dry; related to A. tuberosa, differs by having glabrous stems and narrow
leaflets.
Arachis paraguariensis Chod. et HassI.

Root narrow, elongate, tuberous; stem erect, 20-30 cm tall; the basal
leaflets without lamina or only with two leaflets, rest tetrafoliolate; stipules
15-20 mm long, partially adnate to petiole, free part 10-13 mm long,
linear, softly hairy; petiole 12-25 mm long, woolly; leaflets oblong-
lanceolate, slightly mucronate, marginal vein thickened, midrib with 8-12
secondary veins regularly spread, glabrous above and light green hairy
below, margin woolly, 10-24 x 6-9 mm; flowers mostly subterranean;
hypanthium 14-30 mm long, densely hairy; calyx bilabiate, upper lip four-
toothed, hairy, 5 mm x 2.5 mm, lower lip entire, hairy, 7 mm wide;
standard orange, 16-18 mm wide; wing emarginate; keel pointed, rostrate;
stamens nine, five anthers globose, four elongate; pegs horizontal, may
grow up to 80-90 cm, pods articulate, 8-11 mm x 4-6 mm, with slight
beak, seeds light tan, 5-8 mm x 3-4 mm.
Arachis benthamii Handro

Perennial; roots partially tuberous when old; stem erect, 40 cm or more,
decumbent, simple or branched, terete, hirsute; tetrafoliolate; stipules
narrow-lanceolate, acuminate, veined-striate, hairy outside, glabrous or
sparsely hairy inside, margin ciliate, 25-32 mm long, 8-10 mm adnate to
the petiole; petiole 18-22 mm, grooved, very hairy; rachis 5-6 mm long,
grooved, hairy; leaflets, subcoriaceous, ciliate, basal pair normally obovate
or obovate-oblong, apex obtuse, rounded or slightly emarginate, mucro-
nate, base obtuse, usually hairy above, 32-37 mm x 17-21 mm; hypan-
thium filiform, 6 cm long; calyx deeply bilabiate, upper lip entire, hairy
outside, hairs setaceous, glabrous inside; standard orange, glabrous; pod
uni- or biarticulate, isthmus thread-like, 5 cm long, articles of pod oblong
or obovate, beaked, woolly, hairs later fall off, 8-11 mm x 4-6 mm,
pericarp thin, reticulate; seeds reddish.
Arachis martii Handro

Perennial; stem prostrate, branched, hirsute, 10-17 cm long; tetrafoliolate;
stipules linear-lanceolate, veined-striate, hairy outside, margin cili-
ate, reaching to the tip ofrachis or beyond, 12-15 mm long, 4-5 mm adnate
to the petiole; petiole 8-10 mm long, hirsute, grooved, rachis 2.5-3.5 mm
long; leaflets subsessile, membranaceous, ciliate, basal pair elliptic, apical
pair obovate or suborbicular, tip obtuse or rounded or slightly emarginate,
slightly mucronate, base obtuse, puberulous above, adpressed hairy below,
7-12 cm x 4-9 mm; hypanthium thread-like, hairy, 4 cm long; calyx deeply
bilabiate, upper lip four-toothed, hairy outside, hairs setaceous; standard
glabrous, yellow.
The above description is from a herbarium specimen only. Seed and
germplasm of this species are not available. Several collecting missions to
the recorded area of distribution of this species, including one by the
authors, were in vain.
Arachis rigonii Krap. et Greg.

Perennial, herbaceous; no rhizomes or stolons, lateral roots horizontal;
stems cylindrical, short, 4 cm, glabrous or subglabrous; well-branched,
creeping; tetrafoliolate; stipules falcate, adnate portion 13-15 mm long
and 3 mm wide with rigid spines on outer surface, rest 10-12 mm free,
2.5 mm wide at the base, marked with longitudinal veins, hairy margin;
petiole 6 cm, grooved, rachis 10-15 mm long, grooved; leaflets obovate,
apical leaflets 50 mm x 32 mm, basal 45 mm x 28 mm, glabrous above
hairy below, mucronate; flowers four or five per axil; hypanthium thread-
like, 4-7 cm long, subglabrous; calyx bilabiate, upper lip irregularly three-
or four-toothed, lower lip entire, falcate, standard orange or yellow with
orange markings, 12-14mm x 15-17mm; wings yellow, 8mm x 6mm,
tongued; keel 6-7 mm, beaked, curved, subfalcate; pegs 5-25 cm, hairy at
the base; fruits biarticulate, 11-13mm x 6-7mm, isthmus 1-4cm long,
peri carp slightly reticulated, covered with hairs, prominently beaked;
seeds 9 mm x 4 mm, light tan.
Arachis repens Handro

Perennial; taproot obtriangulate, many secondary roots without thicken-
ings; stem well-branched, often hollow; stem, branches, outer surface of
stipules, petiole, rachis, lower leaf surface and outside of calyx covered
with fine adpressed hairs; stems and branches prostrate or ascending in
shady places, terete, adventitious roots at nodes; internodes 3-5 cm long,
with adpressed hairs; stipules veined-striate, 15-20 mm long, free portion
as long as or shorter, lanceolate or falcate, glabrous or sometimes with few
hairs, adpressed at the base, ciliate margin; petiole 2-4 cm, grooved; rachis
3-7 mm long; leaflets membranaceous, elliptic-oblong, or obovate-
oblong, tip obtuse or rounded, slightly mucronate, base obtuse, glabrous
above, 20-35 mm x 8-12 mm; hypanthium thread-like, hirsute with
short hairs, 8.5-11.0 cm long; calyx deeply bilabiate, 7 mm long, upper lip
three-toothed, lower lip two-toothed, setaceous outside; standard yellow,
8-13 mm x 16-17 mm; wings and keel yellow; fruit biarticulate, peg 5 cm
long, isthmus 1.5-3.5 cm long, articles 8-13 mm x 5-6 mm.
Arachis prostrata Benth.

Perennial; taproot and lateral roots form lomentiform tuberoids, 9 mm x
5 mm; main stem thin, erect, becomes prostrate later on, 5-12 cm long,
covered almost totally by stipules; centrally branched, lateral branches
procumbent, 90 cm long; internodes on main stem short, 5 mm long,
villous, on laterals longer; tetrafoliolate; on main stem stipules rigid,
striate, villous at the base, margins ciliate, hairs 1.5-2.0 mm long, adnate
portion 8 mm long, free portion 10-12 mm and 2 mm wide at the base;
rachis 5 mm long; leaflets commonly obovate, or oblong, 19-20 mm x
11-12 mm; on prostrate laterals stipules shortly adnate, 3-4 mm and free
portion 5-9 mm long and 2.0-2.5 mm wide at the base; petioles 3-5 mm
long, grooved and villous; leaflets obovate, 9-13 mm X 7-9 mm, mucro-
nate, margins hairy, glabrous or slightly hairy above, densely hairy below,
upper surface shining; flowers larger on laterals than on main stem;
hypanthium 2-4 cm, densely hairy; calyx bilabiate, villous, upper lip 4 mm
long, lower lip entire; standard orange with purple markings on the back
(dorsal) face, 8 mm x 9-10 mm; wings yellow, 5 mm long; peg 1-5 cm
long, aerial portion purple and hairy, underground portion with adventi-
tious roots, isthmus short, fruits biarticulate, rarely form a single cavity
with two seeds, 5-8 mm x 4-5 mm, covered with hair; seeds 4-6 mm x
3-4 mm, tan or reddish brown.
Arachis marginata Gard.

Perennial; taproot with many laterals with numerous tuberiform thicken-
ings, 10-25 cm x 5-7 cm; stem erect, 10-20 cm, hardly branched; long and
silky hairs cover the whole plant except the upper leaf surface; internodes
12-30 mm long, quadrangular, villous; tetrafoliolate; adnate portion of
stipules 7 mm long and 5 mm wide, villous and free portion 13 mm long,
glabrous; petioles of basal leaves short, 2 mm long, and apical leaves 3 cm
long, grooved; rachis 7-12 mm long, grooved; leaflets oblong-obovate,
apical leaflets 40-44 mm X 26-30 mm, basal leaflets 32-36 mm x 20-
25 mm, upper surface shining, smooth, glabrous with one line of long hairs
on the midrib; lower surface with thickened veins, reticulate, with long
adpressed hairs, hairs rigid and denser on the midrib, margin thickened,
especially on lower surface, setose, sometimes with short hairs; flowers
small, covered by stipules on basal nodes; hypanthium thread-like, 40 mm
long, villous; standard yellow with purple lines on the back, lines less
distinct on front face; pegs long, up to 60 em, mostly horizontal; fruits
articulate, articles 14-16 mm x 5-8 mm, highly reticulated, strongly
beaked; seeds reddish, 7-10 mm x 4-6 mm.
Arachis villosulicarpa Hoehne
Perennial; long thickened taproot; main stem erect, up to 60 cm, angular,
densely whitish villous, internodes 2-4 cm long; well-branched, branches
prostrate up to 1 m or decumbent; tetrafoliolate; stipules lanceolate, acu-
minate, lower 10-17 mm adnate to petiole, upper 20-30 mm free, 3-9 mm
wide at the base, fine linear margins, sparsely hairy; petiole 19-50 mm
long, grooved, abruptly reflexed below; rachis 5-13 mm long, grooved,
sparsely hairy; leaflets elliptic or oblong-elliptic, apical leaflets 20-50 mm
x 6-22 mm, basal leaflets 16-40 mm x 5-15 mm, marginal veins slightly
thickened, thinly puberulous or glabrous, shiny upper surface, ciliate
margin; flowers axillary, on main stem covered by stipules, one to many in
axils; hypanthium slender, hairy, 2-5 cm long; calyx bilabiate, upper lip
entire or two-toothed, lower lip thin, linear, 5-6 mm long, hairy outside;
standard suborbicular, orange with purple markings on the back, reflexed,
10-12 mm X 12-14 mm; wings oblong, orange with yellow blush at the tip;
keel incurved, beaked; fruit biarticulate, pegs 6-26 cm, pod exterior
densely villous or covered with tomentose hairs, 20-25 mm x 7-9 mm;
seeds oblong, yellow-purplish, 16-23 mm x 6-7 mm.
A. villosuficarpa is considered as a cultigen in the genus Arachis, culti-
vated by a small group of Amerindians in north-western Mato Grosso state
of Brazil. This species has not been found in the wild (Valls et af., 1985).
Interspecific hybrids with A. hypogaea have been reported (Raman, 1976;
Sundaram, 1985), however similar attempts at hybridization elsewhere
have failed (Smartt, 1990).
Arachis lutescens Krap. et Rig.

Perennial without rhizomes or stolons; root with thick and elongated
tubers about 10 cm x 5 cm; most of the aerial parts covered with hairs
except the upper leaf surface; stem erect, 3-10 cm tall, laterals procum-
bent, 60 cm long; internodes 10-25 mm long; tetrafoliolate, rarely with a
supernumerary apical leaflet; on main stem adnate portion of stipule
subfalcate, 6-8 mm x 2 mm, free portion linear-Ianceolate, 10-16 mm
long; petioles on main stem longer, 15-45 mm long, shorter on branches,
2-6 mm long; leaflets on main stem oblong, sometimes obovate, apical
leaflets 15-17 mm x 9-12 mm, basal leaflets 21-23 mm x 8-10 mm; obo-
vate or oblong on branches, apical leaflets 8-18 mm x 6-11 mm, and basal
leaflets 15-16 mm x 9-11 mm, upper leaf surface glabrous, shining, lower
surface hairy, hairs 2 mm long, sometimes adpressed; hypanthium thread-
like, villous, 2-4 cm long; calyx villous, bilabiate, upper lip irregularly
three-toothed, lower lip entire, linear-acuminate, subfalcate, 6 mm long;
standard orange with central yellow portion, suborbicular, 10 mm x
11 mm; wings 6 mm long, yellow with orange blush in front; keel falcate;
fruits biarticulate, pegs 2-5 cm long, aerial portion pubescent, under-
ground portion without roots, articles of pods 6-9 mm x 5-6 mm, apical
article generally longer, pods densely hairy, isthmus short, 1-3 mm long;
seeds 5 mm x 3.5 mm.
Arachis burkartii Handro

Perennial; rhizomatous, rhizomes 35 cm x 2 cm; stem short, erect, simple
with little branching, branches short; stipules narrow-Ianceolate, long-
acuminate, striate, hairy outside, ciliate, 15 mm or longer, one third adnate
to the petiole; petioles long, 7 cm, terete, slightly grooved above, hairy;
rachis 4-8 mm long; leaflets rigid, coriaceous when dry, ciliate, subsessile,
elliptic or obovate or obovate-Ianceolate or obovate-oblong, apex acute,
mucronate; base obtuse or rounded, glabrous above, adpressed hairy
below, thick-nerved, 2-41 cm x 10-12 mm; hypanthium thread-like, hairy,
6-9 cm long; calyx deeply bilabiate, upper lip entire, outside hairy; hairs
setaceous, sparsely mixed; calyx hairy, bilabiate, upper lip four-toothed,
lower entire and acuminate; standard suborbicular, orange; wings oblong,
obtuse; pods 10 mm long, peri carp slightly reticulated; seed tan, tip
elongated or rounded.
Arachis glabrata Benth.

Perennial; rhizomatous, prostrate or ascending; stems and rhizomes thick,
rhizomes subterranean, long, branched; stem semi-woody, short, well-
branched; branches prostrate, subglabrous; stipules narrow, acuminate,
20 mm x 2 mm; petiole 3-6 cm long, sparsely hairy, hairs long; tetrafolio-
late, leaflets oblong-elliptic, 22-30 mm x 10-18 mm, tip acute or some-
times obtuse, glabrous above, rarely hairy below, margins may be
serrulate; numerous flowers but few fruits; standard yellow or orange-
yellow, 15-20 mm wide; hypanthium 3-6 cm long, hairy; calyx hairy,
bilabiate, upper lip four-toothed, lower entire and acuminate; standard
suborbicular; wings oblong, obtuse; pods 10 mm long, pericarp slightly
reticulated, seed tan, tip elongated or rounded.
Wide variation for morphological features occurs between accessions of
A. glabrata collected so far. Variation in plant size, growth habit, leaf
canopy, leaflet shape, size and hairiness and flower colour have been
observed. Seed set is rare though not uncommon in the area of
distribution.
Arachis hagenbeckii Harms.

Perennial, central taproot with rhizomes; rhizomes long, 10-35 cm, well-
branched, 1.5-5 mm thick with adventitious roots; stems prostrate, subgla-
brous; tetrafoliolate; stipules 30 mm x 4-7 mm; petiole 4-8 mm long,
glabrous or sparsely puberulous, at the point of attachment to the leaflet
densely hairy; leaflets oblong, elliptic-lanceolate, narrowly oblong or
obovate-oblong, base obtuse or rounded, tip obtuse to subacute, slightly
mucronate, glabrous on both surfaces, subcoriaceous, marginate, 30 mm x
8-15 mm; long hypanthium, subglabrous or puberulous, 5-7 cm long; calyx
Infraspecific differentiation and morphology of 53
falcate, bilabiate, upper lip irregularly four-toothed, lower lip entire, up to
10 mm long, sparsely hairy outside; standard suborbicular, 12-15 mm
wide, yellow; keel hyaline, falcate; style hairy at the tip; pegs 6 cm long;
fruits rare, generally uniarticulate, ovoid, acuminate, 16 mm x 8 mm.
Arachis pusilla Benth.

Perennial (annual?); taproot well-branched; main stem short, 3-8 cm tall,
erect, angular, densely hairy; basal branches thin, procumbent or decum-
bent, subglabrous; tetrafoliolate; on main stem adnate portion of stipules
10-12 mm long, free portion 20-23 mm, minutely puberulous at the mar-
gin, hairy at the base, petioles 50-60 mm long, rachis 17-19 mm; on lateral
branches adnate portion of stipules 5-7 mm, free portion 10-17 mm,
petioles 25-40 mm long, rachis 8-15 mm long; leaflets ovate or obovate on
main stem; apical leaflets 29 mm x 17 mm, basal 28 mm X 16 mm, leaflets
on branches oblong-obovate, apical leaflets 17-26 mm x 9-17 mm, basal
leaflets 15-26 mm x 14-17 mm, membranaceous, acute, glabrous or
adpressed pilose, mucronate; hypanthium thread-like, 37-43 mm long,
slightly pubescent; calyx bilabiate, upper lip irregularly three- or four-
toothed, lower lip entire, hairy outside, 5 mm long; standard orange,
crescent yellow, suborbicular, 10 mm in diameter, wings lemon yellow,
6-7 mm long; keel hyaline, falcate; fruit 2-3 articulate, articles 10-12 mm
x 6-7 mm, covered with short hairs, pegs almost horizontal, 30-72 cm
long, inserted basally to pod, isthmus 20-37 cm long; seeds 8 mm x 4 mm,
light tan.
The name A. pusilla for the above description may be incorrect (Krapo-
vickas, 1988) but it is given here because there is as yet (1994) no authentic
renaming of the species.
3.4 INFRASPECIFIC DIFFERENTIATION AND MORPHOLOGY

OF ARACHIS HYPOGAEA
As in the case of interspecific taxonomy of the genus Arachis, subspecific

classification of A. hypogaea has been considered by various workers (for a
review see Ramanatha Rao, 1987; 1988). Most of the earlier classifications
were based on growth habit, presence or absence of dormancy and matur-
ity. The later attempts included characters such as branching pattern and
location of reproductive branches. The classification by Gregory et al.
(1951) was a comprehensive study in which groundnut was divided into two
large botanical groups, virginia and spanish-valencia, on the basis of
branching pattern as described by Richter (1899). The presence or absence
ofreproductive axes (inflorescence) on the main stem and the arrangement
of reproductive (R) and vegetative (V) axes on the primary laterals were
the most important criteria. The main axis was denoted as n, and the
primary, secondary, and tertiary lateral branches 0+1, 0+2, and 0+3,
respectively.
The virginia group is characterized by the absence of R axes on the main
stem. Further, alternating pairs of V and R axes are borne on the cotyle-
donary lateral and other 0+1 branches. This system is termed the 'alternate
branching pattern'. The first two branches on the 0+1 lateral are always
vegetative. The alternate branching pattern is repeated in higher order
branches, with some irregularity, especially on the distal nodes. The
spanish-valencia group is characterized by the presence of R axes in a
continuous series on successive nodes of lateral branches, on which the first
branch is always reproductive. This system is called the 'sequential branch-
ing pattern'. R axes are also borne directly on the main axis at higher
nodes. Most 0+2 and 0+3 nodes are reproductive. Generally the spanish
types show some amount of 0+2 branching in an irregular fashion.
Valencia types may not have any 0+2s, if present they will be in a
sequence, distal to the 5th to 8th nodes of the 0+ 1 branch.
A few other systems of classifications were also attempted based on
morphological characters, with different degrees of importance given to
characters such as growth habit and branching pattern (Bhavanisankar Rao
and Raman, 1960; John et al., 1954; Seshadri, 1962; Varisai Muhammad et
al., 1973a; Varisai Muhammad et at., 1973c). All these later systems are
deficient because either the germplasm considered for classification was
limited or the systems did not take into account the classical taxonomic
nomenclature (Dubard, 1906; Harz, 1885; Waldron, 1919). Krapovickas
studied extensive collections and proposed a system of subspecific
classification of A. hypogaea (Krapovickas, 1969; Krapovickas, 1973;
Krapovickas and Rigoni, 1960). In this system, subspecies were found to
be associated with the five geographic regions: Guaranian, Bolivian,
Peruvian, Amazonian (Rondonia and north-west Mato Grosso) and the
region of Goias and Minas Gerais. The number of regions was extended to
six, to include north-east Brazil (Gregory and Gregory, 1976). This com-
prehensive subspecific classification is summarized below, along with a
brief description of each class:
Arachis hypogaea L. (Linnaeus, 1753)

Subsp. hypogaea (Krapovickas and Rigoni, 1960)
Habit procumbent, decumbent or erect; branching alternate; inflorescence
simple and never borne directly on the main axis, first branch on the
cotyledonary lateral always vegetative; two or two to four seeds per pod;
pod beak prominent, slight or absent; pod constriction prominent, slight or
absent; pod very large (20 mm) or small «10 mm); generally tan seed coat
colour but red, white, purple and variegated forms exist; seed dormancy
usually present; foliage dark green.
Var. hypogaea (= A. africana Lour; type 'Braziliano' Dubard, 1906;
type 'Virginia' (Gregory et al., 1951)); Bolivia and Amazonia. Habit
Morphology and development 55
procumbent, decumbent or erect; main aXIs In procumbent forms
short (not exceeding 40-50 cm); stem usually not very hairy; usually
two-seeded; pod beak generally not very prominent, medium-late
maturing.
Var. hirsuta Kohler (= A. asiatica Lour; type 'peruano' (Dubard, 1906
in part)); Peru. Habit procumbent; main axis may exceed 1 m; stem
fairly hairy; pods strongly beaked and ridged, with two to four seeds;
very late maturing.
Subsp. Jastigiata Waldron (Waldron, 1919)
Habit erect or decumbent; branching sequential; inflorescence simple or
compound, always present on main axis; first branches on cotyledonary
laterals reproductive; seed dormancy usually absent; foliage usually lighter
in colour than in subsp. hypogaea
Var. Jastigiata (= type 'peruano' (Dubard, 1906 in part)); type 'valencia'
(Gregory et al., 1951); Guarania, Goias, Minas Gerais, Peru, and
north-east Brazil. Vegetative branches on primaries absent or regu-
larly placed at the distal nodes; inflorescence usually simple; pods with
two or two to four (rarely five) seeds; beak absent, slight or promi-
nent; size medium to small; testa colour tan, red, white, yellow,
purple or variegated.
Var. vulgaris Harz (= type 'spanish' (Gregory et al., 1951)); Guarania,
Goias, Minas Gerais, and north-east Brazil. Vegetative branches
occasional and irregularly placed; inflorescence compound; pods
usually with 2 seeds; beak present or absent; size medium to small;
testa colour tan, red, white, or purple.
There have been some attempts to link the morpho-agronomic classifi-
cation (Bunting, 1955; 1958; extended by Smartt, 1961) with the taxonomic
treatment (Krapovickas, 1969; Krapovickas and Rigoni, 1960) by some
workers (Gibbons et al., 1972; Varisai Muhammad et al., 1973a, 1973c).
These attempts were based on studies of limited amount of germplasm
available to the authors. However, much larger geographically and geneti-
cally more diverse collections cannot be explained by any of the above
systems. With the many intermediate forms that are now available, any
classification will be very difficult and may not be of great practical value
except locally.
3.5 MORPHOLOGY AND DEVELOPMENT
The morphological description and developmental aspects of groundnut

were very confused for a long time. The geocarpic fruit, the complex
branching pattern and the highly condensed (telescoped) nature of the
reproductive axis (inflorescence) have been mainly responsible for such a
confusion. Many of the earlier workers were unable to associate the
underground fruits to the plant or to the aerial flower. Marggraff (1648)
illustrated the fruits as growing on the roots. The first accurate description
of flowers was published by Poiteau (1806). However, it was only in 1950
that Smith published for the first time a clear and correct account of the
aerial flowers and subterranean fruits of the groundnut (Smith, 1950). A
brief, updated description was published by Hammons (1981).
In the following account the developmental aspects are combined with
morphological description. This is because morphology at any point in time
is best described in relation to the developmental stage.
3.5.1 Seed
Seeds show large variation for their size, shape and colour. The testa or
seed coat is thin and papery. Generally the seed coat constitutes three
unicellular layers: the epidermis or sclerenchyma, the middle parenchyma
and the inner parenchyma. These layers represent the integuments of the
maturing ovule and are maternal in origin (Glueck et al., 1979). The
surface as well as the transverse sections of testa show a great deal of
diversity (Zambettakis and Bockellee-Morvan, 1976). Groundnut cultivars
can be grouped according to the size of the wax layer, the joining of the
epidermal cells, thickness of cell walls and presence of cracks in the
epidermal layer. The seeds of wild species basically resemble the cultivated
groundnut, except that they are much smaller in size.
(a) Size
Seed size is an important economic character. It is fairly stable for any
given cultivar and is highly diagnostic in nature. Seed lengths ranging from
7 to 21 mm and seed diameters from 5 to 13 mm have been observed
(Ramanatha Rao, 1988; Retamal et al., 1990). Seed size, together with the
seed mass, has been used extensively in agronomic classification of ground-
nut. Larger seed types are preferred for confectionery purposes, while
most of the oil types have medium to small seeds. Seed size in wild Arachis
is much smaller and length ranges between 8 and 18 mm (16-23 mm in
A. villosulicarpa) and diameter between 4 and 7 mm.
(b) Mass
Seed mass (weight) is an important economic as well as diagnostic charac-
ter. Depending on the material studied and site of evaluation, various
ranges have been reported: 0.2-1.0 g (Seshadri, 1962); 0.17-1.24 g (Rama-
natha Rao, 1988); and 0.54-2.38 g (Retamal et al., 1990). In general,
cultivars belonging to var. hypogaea tend to have larger and heavier seeds;
those belonging to var. Jastigiata have smaller and lighter seeds; and the
wild species have much lower seed mass.
(c) Colour
Colour of the seed coat or testa is an important diagnostic character. It is
also an important market trait. Broadly, groundnuts can be classified into
those possessing non-variegated testa (one solid colour) and those having
variegated testa (more than one colour). Generally, seed colour deepens
with storage in shell or as shelled seed over a period of time (Bunting,
1955) so that observations immediately after harvest or a long time after
harvest can be unreliable. Different solid colours such as white, rose,
flesh, wine, red, light purple and dark purple can easily be recognized.
Within each class, the intensity may vary depending on maturity, environ-
ment, genotype or the interaction between genotype and environment.
Variegated testa colour has been associated with splitting or rupture of the
outer epidermis due to differential growth rates of testa and embryo (Ashri
and Yona, 1965; Stokes and Hull, 1930; Yona, 1964) and inhibition of full
development of the outer epidermal layer of testa in some regions (Branch
and Hammons, 1979). However, these theories can explain only the
variegation when white is involved. The exact genetic or physiological
mechanisms for variegation of testa are yet to be fully understood. In some
cases a white spot may appear on the seed coat opposite to the micropylar
end (Srivastava, 1968).
The various solid colours observed in cultivars include off-white, yellow,
pale tan, light tan, dark tan, rose, grey-orange, light red, red, dark red,
light purple and dark purple. Among the variegated types, two com-
ponents can be recognized: the major (primary) colour and the minor
(secondary) colour superimposed on the major colour. The four observed
colour combinations are: red and white, purple and white, light tan and
dark tan, tan and purple. In the case of combinations involving white, it
can be almost pure white or off-white. Similarly, different shades of purple
and red occur. The minor colour may vary in its degree from thin streaks to
large blotches. Because of the subjective nature of this character, use of
guides such as the Royal Horticultural Society (RHS) Colour Chart is
recommended (IBPGRJICRISAT, 1981). Seed colour is best observed at
least two weeks after harvest (not later than six weeks) and after thorough
drying.
Wild Arachis species show much less variation. The main colours that
have been recorded in different species are: yellowish-tan, light tan, dark
tan and reddish brown.
(d) Primordia in seed

Each seed consists of two massive cotyledons, upper stem axis and young
leaf primordia (epicotyl), hypocotyl and primary root. Unlike other seeds
of Papilinoideae, the embryo of the groundnut is straight. It contains all
the primordial leaves and the above-ground parts that appear during the
first two weeks of growth. The epicotyl consists of three buds, one terminal
with four leaf primordia and two cotyledonary laterals with one or two leaf
primordia (Gregory et al., 1951; Maeda, 1972). Some instances of poly-
embryony have been recorded (Patel and Narayana, 1935; Raman and
Nagaraj an , 1958; Smith, 1950). The speculation about the presence of
flower primordia in seed (Schwabe, 1971) could not be confirmed (Star-
itsky, 1973).
3.5.2. Seedling
The morphology, growth and development of the groundnut seedling was
studied by a number of workers (Badami, 1933, 1935; Bouffil, 1947;
Richter, 1899; Yarbrough, 1949, 1957a, 1957b). The radicle consists of
about half hypocotyl and half primary root during the first few hours of
germination, depending on the depth of planting. In moist soils at 27°C,
the primary root emerges in 24-36 hr. The primary root may grow from 0.5
to 4.0 cm in four days. Lateral roots generally appear after the second day,
and as many as 100 laterals could be formed in five days (Yarbrough,
1949).
3.5.3 Root system

Groundnut is a herbaceous annual with a taproot and a fairly well-
developed root system (Figure 3.1). However, the root and hypocotyl may
be modified to form tubers (section Erectoides) or tuberoids (sections
Erectoides and Extranervosae). Diameter of the primary root may vary
from a few millimetres in annual species to 8 cm in perennial species
(Gregory et al., 1980). Adventitious rooting at nodes of branches that
come in contact with soil is common, and is more frequent in species
belonging to section Caulorhizae. Most of the root system is generally
concentrated at a depth of 5-35 cm (Narasinga Rao, 1936). The root
spread is confined to a radius 12-14 cm. The spreading types usually have a
more vigorous root system than the bunch types (Mohammad and Khanna,
1932; Seshadri et al., 1958).
(a) Taproot
On the second day after germination the taproot appears with a large root
cap. It elongates rapidly and grows almost vertically downwards, diverting
only to overcome obstacles. The taproot can grow to a length of 50-55 cm
in ill-drained clayey soils (Badami, 1935), or up to 90-130 cm (Bruner,
1932). The root elongation depends on the cuitivar, soil and available
j---Leaflet
Stem
Flower
Peg
Young Pod
Mature Pod
Tap Root
Figure 3.1 Groundnut plant.
moisture (Intorzato and Tella, 1960; Seshadri et al., 1958; Yarbrough,

1949).
(b) Lateral roots

Lateral roots appear on the third day after seed germination in the upper
region of the taproot. As many as 100-120 may be produced by the fifth
day which grow to a length of 15-20 cm (Yarbrough, 1949). The lateral
roots are basically similar to taproots. The main differences are the lack of
central pith and being diarch in contrast to the tetrarch primary root
(Gregory et al., 1973).
(c) Hypocotyl
The hypocotyl (the portion of the stem which lies between the primary root
and the cotyledons) is succulent early on and grows up to 10 mm in
diameter. The region between the hypocotyl and the primary root is
bounded by the intact epidermis of the hypocotyl (Yarbrough, 1949). This
collar region is swollen due to the enlargement of the cortex. It elongates
to bring the cotyledons to the soil surface and its length depends mainly on
the depth of planting (Bouffil, 1947). The emergence is complete in about
10 days after sowing and the hypocotyl grows to 1-11 cm (Bouffil, 1947;
Seshadri, 1962; Yarbrough, 1949). It loses its fleshy nature, becomes
fibrous and changes from white to creamy-white to brown, becoming
indistinguishable from the taproot by about four weeks after germination.
Later it may produce many roots.
(d) Root hairs
The outer layers of the primary root generally slough off as root growth
proceeds (Petit, 1895) causing loss of the structural base for root hairs
(Richter, 1899). However, under certain conditions formation of root
hairs, as clusters or rosettes in the junction of root axils, has been observed
(Badami, 1935; Reed, 1924; Waldron, 1919). These hairs were less fragile
than those found in other legumes and ranged from 2 mm to 4 mm in length
(Allen and Allen, 1940). Some of these hairs were septate (Chandler,
1978). These structures have often been referred to as root hairs, a
misnomer in the strict botanical sense.
Though root nodules are not plant structures in a botanical sense, their
close association with the plant makes them appear so. Nodules first
appear when plants are about 15 days old. A group of nitrogen-fixing
bacteria belonging to the genus Bradyrhizobium is responsible for the
formation of nodules. Though groundnut does not possess real root hairs,
an association between the tufts of hairs in root axils and nodulation has
been found (Nambiar et at., 1983 ).
3.5.4 Stem
The stem is generally angular, pubescent and solid with a large central pith
in the early stages (section 3.6.2). As the plant grows, stems tend to
become hollow and cylindrical and shed most of the hair, especially on
lower internodes. There is no indication of woody development in A.
hypogaea but some wild species can develop woody stems (section 3.3).
The main stem develops from the terminal bud of the epicotyl flanked by
two opposite cotyledonary laterals. Though main stem height depends
on genotype, it is influenced by environment to a considerable degree and
ranges from 12 cm to 65 cm (Ramanatha Rao, 1988). In wild Arachis, the
main stem tends to be much shorter, ranging from 12 cm to 35 cm, with a
few exceptions. Stem thickness is highly variable, although Seshadri (1962)
reported that generally the bunch types have thicker internodes, short and
highly condensed at the base and longer at the higher nodes. The basal
stem diameter could be as much as 8 cm in some of the wild species.
(a) Colour
Anthocyanin pigments in the epidermal cells of the stem can give different
shades of colour. Stem colour is determined by the absence or presence
and intensity of pigmentation. The common colours observed are purple,
pink, dark red, light red or green (absence of anthocyanin) and numerous
shades of purple, pink and red. The colour development is influenced
significantly by exposure to sunlight and recording typical colours is gener-
ally difficult. Hence, the stem colour could be recorded as present or
absent, classifying the cultivars into two broad groups (IBPGRlICRISAT,
1981).
(b) Hairiness
Groundnut has been described as a glabrous to hirsute herb, indicating the
extent of variability for this character. A range of two to four grades of
hairiness were recognized (Hayes, 1933; John et al., 1954; Patel et al.,
1936; Patil, 1965; Varisai Muhammad et al., 1973c). Because of the diffi-
culty in defining various grades of hairiness, it is pragmatic to recognize
only two grades: scarce and abundant. Generally the upper internodes
should be observed because at lower levels the hairs are not persistent. The
hairs are arranged in regular rows on the stem. It is possible, by careful
observation, to classify the arrangement into the number of rows of hairs
(two, four, six and irregular).
Usually the following three types of hairs have been observed on plant
parts:
• Long hairs (up to 3 mm), are generally septate and uniseriate and
distributed irregularly. Occasionally the cell walls of the top three or
four cells disintegrate, giving the appearance of a single long cell. The
outer walls may sometimes be thickened.
• Short hairs «1 mm), generally occur densely along with the long hairs.
• Glandular hairs (trichomes, spines) are long and bristle-like, green
with chlorophyll and multicellular with a bulbous base.
(c) Growth habit

The groundnut plant has a distinct main stem and a variable number
of lateral branches. The carriage of laterals determines the growth habit of
the plant. Two distinct forms of growth habit - spreading (runner, trailing,
procumbent and prostrate) and erect (upright, erect bunch and bunch) -
have long been recognized and have provided the basis for both agronomic
and taxonomic classifications.
There has been no agreement on classification of growth habit in ground-
nut (for a review, see Ramanatha Rao, 1988). This is further complicated
by the complexity of inheritance of this character (Hammons, 1973; Wynne
and Coffelt, 1982). A large number of intermediate forms occur because of
cytoplasmic and genetic factors that interact together and also interact with
light environments for the expression of growth habit (Ashri and Goldin,
1963; Ziv et al., 1973). Recognizing bunch (erect) and runner (trailing)
forms as the major forms, it was suggested that the cytoplasmic-genic
interactions that determine growth habit act on the biosynthetic pathways,
producing phytohormones and their inhibitors (Ashri, 1976; Resslar and
Emery, 1978).
The above discussion suggests that growth habit is not a discrete charac-
ter, and its inheritance is complex. Several loci interacting among them-
selves and with the cytoplasm may produce various grades. However, for a
given homozygous accession, its growth habit can be defined as stable.
In the earlier classifications, the runner and spreading bunch forms
were associated with alternate branching (subsp. hypogaea) and the erect
habit with sequential branching (subsp. fastigiata) (Krapovickas, 1969;
Krapovickas, 1973). However, if one considers the enormous variation
that exists in the current groundnut genetic resources, such association is
no longer very clear-cut. Hence, the need to reclassify the growth-habit
types is recognized. Growth habit needs to be described purely on the basis
of observations on plants grown at a wide spacing in a given environment.
Nevertheless, it must be noted that a vast majority of runner and spreading
bunch forms show alternate branching, while the erect forms are generally
sequentially branched. The major collections would include, along with
landraces, a number of accessions derived through breeding and also
accessions from South America. A certain amount of introgression might
have occurred at subspecific level in the centre of diversity. In such
germ plasm the following growth habits could be observed which are based
only on the carriage of plant and the position of primary branches in
relation to the main stem (IBPGR/ICRISAT, 1981) (Figure 3.2):
1. Procumbent 1: The main stem is erect and may vary in height. The
lateral branches are prostrate.
2. Procumbent 2: This is similar to Procumbent 1, but the main stem has
~~.
1 Procumbent - 1 2 Procumbent - 2
3 Decumbent - 1 4 Decumbent· 2
5 Decumbent - 3 6 Erect
Figure 3.2 Growth habit.

a tendency to bend and continue to trail on the ground. This type is
generally encountered in wild Arachis.
The common runner (= spreading, trailing, creeping or prostrate)
habit group includes the above two forms.
3. Decumbent 1: The main stem is distinct. The laterals, which normally
trail on the ground, tend to become upright at distal nodes.
4. Decumbent 2: The main stem is distinct. The laterals tend to be almost
upright at the median nodes.
5. Decumbent 3: The main stem becomes indistinct as most of the lateral
branches are almost upright and give the plant a bushy appearance.
The spreading bunch (= semi-spreading, bunch runner, and runner
bunch) group generally includes the above three forms.
6. Erect: the main stem is indistinguishable from the laterals. The laterals
are at an acute angle to the main stem. This form includes the erect
bunch or bunch groups in the earlier classifications.
(d) Branching pattern

The arrangement of reproductive (R) axes on the main stem (D) and Rand
V axes on the primary lateral branches (0+ 1) is the basis for classifying the
branching pattern. Groundnuts were classified into two botanical groups
based on the branching pattern (Gregory et at., 1951), which were named
as alternate branching and sequential branching (Bunting, 1955) (section
3.4). Such a system was confirmed and used in classifying A. hypogaea
along with other correlated characteristics (Krapovickas, 1969). The culti-
vars of var. hypogaea produce numerous secondary (0+2) and tertiary
(0+3) branches, while subsp. fastigiata produces a limited number of 0+2s
and rarely 0+3 branches.
Describing groundnuts as alternate and sequential forms had been found
adequate until recently. The order of occurrence of vegetative and repro-
ductive nodes on the basal primary branch of A. hypogaea subsp. hypogaea
var. hypogaea was studied. Wide variation was observed in vegetative and
reproductive node numbers and it was suggested that the sequence in
runner types could be a result of introgression. Because of the large
variation, it was felt that branching pattern might not be a dependable
attribute to classify groundnuts (Bhagat et at., 1987). However, more
studies of this nature, over different environments, would be required to
draw such definitive conclusions on such an important character. It is also
essential to note that branching pattern should be observed on the cotyle-
donary lateral, which may be difficult to identify in well-grown runner
types.
Intermediate types from infraspecific crosses have been observed and
there is recognition of the serious problem that such forms can cause in
taxonomic treatment, due to the increased frequency of introgressed
forms, arising from artificial or natural hybridization between the two
1 Alternate 2 Sequential
3 Irregular with flowers 4 Irregular without flowers on main stem

on main stem
o Reproductive branch
Figure 3.3 Branching pattern.
subspecies (Maeda, 1973) . This is indeed so, as the assembly of a large

number of cultivars includes many variants that cannot be assigned to
either group (Ramanatha Rao, 1988). To account for such types it was
proposed to have two more groups: irregular branching-1 (this is similar to
the alternate pattern but R axes occur on the main stem also) and irregular
branching-2 (similar to the sequential pattern but with no R axes on the
main stem) (Figure 3.3).
3.5.5 Leaf
The leaves that first appear are from the primordia in the seed and new
ones are formed much later. Leaves are tetrafoliolate except in species
belonging to the section Trifoliolatae (= series Trifoliolatae of the section
Erectoides), which have three leaflets. Occasionally small and abnormal
leaflets may appear. The leaves, which are paripinnate, are borne spirally
in a 2/5 phyllotaxy and their arrangement on the main stem and higher-
order branches is distichous. Leaflets are opposite, subsessile, elliptic
(variable) and shortly mucronate with entire ciliate margin. Stipules are
prominent, linear and adnate to some length to the petiole and become
free at the pulvinus. Foliaceous stipules have been reported (Mouli and
Kale, 1982). The leaflets are borne on a slender, grooved and jointed
rachis. The leaf exhibits nyctiotropic (sleep) movements, where the adaxial
surfaces of the leaflets come together and the petiole droops during each
night. Considerable variation exists in leaf characters such as foliage
colour, shape, hairiness and size.
Stomata are present on both the leaf surfaces (Smartt, 1976b). Stomatal
frequency and distribution on leaf epidermis were studied and a higher
stomatal count was recorded on the upper surface than on the lower (Lin et
at., 1986).
(a) Colour
Leaf colour is an important character to differentiate the subspecies in
groundnut. A. hypogaea subsp. hypogaea has dark green foliage in contrast
to the light green of subsp. fastigiata. Leaf colour recordings are highly
subjective: differentiating subtle shades always poses problems and the
intensity of light at the time of observation can also create difficulties. The
use of a guide such as the RHS Colour Chart has been recommended
(section 3.5.1 (c)). Some of the wild species, especially those belonging to
section Extranervosae and a few cultivars belonging to var. fastigiata, often
have shiny upper leaf surfaces.
(b) Shape
The shape of leaflets varies from oblong to lanceolate. The variation for
leaflet shape is much greater in wild Arachis than in the cultivated species.
Linear-lanceolate or narrow-lanceolate leaflets have been recorded
(Bhide and Desale, 1970). Various shapes observed include cuneate,
obcuneate, elliptic and lanceolate. However, variation exists for leaflet
shape even within a plant. This variation may be between the leaflets on
basal nodes and the ones on upper nodes and also between the apical
leaflets and the basal leaflets of the same leaf. In most of the wild species,
the shape of the leaflets on the main stem leaves may differ from that of the
leaflets on lateral branches. For standardization purposes, the shape is best
recorded from the fully expanded apical leaflet of the third leaf on the main
stem and also the shape of the apical leaflet of the fifth leaf on primary
lateral in the case of wild species (IBPGRlICRISAT, 1981).
(c) Size
Leaflet size is variable among cultivars, within a cultivar, or even on the
same plant, depending on the position of the leaf on the plant. The apical
pair is generally larger than the basal. Leaflets on the main stem are slightly
larger than those on the branches in both cultivated and wild species. Size
is influenced to some extent by environment. For standardization pur-
poses, the size of the apical leaflet of the third leaf on the main stem is
considered and it is a diagnostic character. Subsp. fastigiata usually has
larger leaflets than subsp. hypogaea. The size of a leaflet varies from 24 mm
X 8 mm to 86 mm x 41 mm. In wild species, the range is much larger:
15 mm x 8 mm to 95 mm x 30 mm (Ramanatha Rao, 1988).
(c) Hairiness
The leaflets are pubescent, as are the stems, mainly on the abaxial surface
and on the margins. Generally midribs are also hairy, often more densely
than the leaf surface. As described earlier (section 3.5.4 (b», all the three
hair types -long, short and glandular - can be found on the petiole, rachis
and leaflet. There is significant variation in hair distribution, shape, size
and number in A. hypogaea. In wild species, stem hairiness (type and
distribution) has taxonomic significance. The length, shape and number of
hairs appear to be associated with resistance to leaf hoppers (Campbell et
al., 1976). Five different grades of hairiness can be distinguished: almost
glabrous, sparse and short, sparse and long, profuse and short, and profuse
and long. Differences in grades have been noticed between young and
mature leaves.
3.5.6 Inflorescence
The inflorescence, which is a reduced simple or compound monopodium,
appears as a cluster of flowers in the leaf axil (Figure 3.4). In some wild
species it appears as a cluster around the crown of the plant, with subterra-
nean ovaries and the hypanthium exposing other flower parts. The inflor-
escence, consisting of up to three (but occasionally more) flowers, is spike-
like and always occurs in the axils of cataphylls or foliage leaves. The
reproductive (R) and vegetative (V) axes appear to occur at the same
nodes because of the short internodes below the first cataphyll branch
(Norden, 1980). The inflorescence is a small replica of the vegetative
branch. In the axils of scale-like leaves, either a very short peduncle
bearing a single flower (simple inflorescence) or a peduncle bearing sec-
ondary branches (compound inflorescence) is produced. In var. hypogaea
the inflorescence expands very little during maturation. In var. vulgaris it
expands moderately, while in var. fastigiata the inflorescence internodes
may elongate to form a conspicuous branch that may terminate with a pair
of small leaflets. This branch may also bear inflorescence and may be
confused with normal V axes. But R axes can be identified by their
numerous scale leaves at the basal nodes. The length of the reproductive
axes is cultivar-dependent and may exceed 10 cm in some cases.
3.5.7 Flowers and flowering

The typical papilionoid corolla is inserted on the top of the hypanthium
and surrounds the staminal column (Figure 3.5). Flowers are enclosed
between two bracts, one of which is simple, subtending a short peduncle,
and the other bifid, subtending the pedicel. The flower is sessile but
appears stalked after the growth of a tubular hypanthium just before
anthesis. The calyx is five-toothed, one of which is free and anterior to the
keel. The length of the posterior lobe in relation to the standard petal
varies from half the height of the standard to three-fourths. The other four
are fused up to near their tips at the back of the standard and sometimes
the calyx is considered to be two-toothed. The colour of the standard petal
is generally orange, with a basal central area, the crescent, which may be
marked with a different colour. The wings are generally yellow, occasion-
ally marked with a diffuse band of deeper yellow, orange or brick-red at
the tip, generally referred to as 'blush'. The keel is almost hyaline; it is
faintly yellow and closely encloses the staminal column. Generally, the
keel and staminal column are curved at right angles (reflexed) in the mid-
section. Under normal conditions the flower stalk (actually the hypan-
thium) is about 4-5 cm long; the standard is 10-22 mm wide, and 8-16 mm
long.
The stamens are 10, monadelphous, with the staminal column surround-
ing the ovary. Two are usually sterile. Of the remaining eight, four are
globose and uniloculate alternating with four oblong anthers, of which
three are biloculate and the one opposite the standard is uniloculate
(Smith, 1950). However, it has been observed that the oblong anthers
become biloculate and globose uniloculate when the anthers reach
maturity (Xi, 1991). The filaments are fused for two thirds of their length
and form acute angles with their fused bases, because at the point of
separation the filaments are sharply bent. Three basic morphological
groups were identified in pollen: spheroidal, prolate and columnar-
spheroidal. Differences among cultivars in shape, size, exine sculpturing
and poration of pollen grains may be useful for varietal classification and
taxonomic studies (Pen et al., 1987). The pistil consists of a single ovary
surrounded by the base of the hypanthium. The ovary is about 1.5 mm long
Flower---
Hypanthium
('Calyx tube') - _ .
Stipule ----l!i'lm~
Figure 3.4 Inflorescence.

Fleshy
staminal
tube
Lower lip of the calyx

Style Keel
Hypanthium (,Calyx tube')
1 em I
Figure 3.5 Flower - a longitudinal section (after Smith, 1950).
and 0.5 mm wide, and has two to four, sometimes five and rarely six
ovules. The long filiform style has two sharp bends and is covered with
upward-slanting hairs near the top on the surface facing the standard
(Gregory et al., 1951; Smartt, 1976a; Omen, 1933). The stigma is club-
shaped or clavate, usually at anther level or protruding slightly above.
(a) Standard petal colour

The most common colour of the standard is orange but other colours such
as white, lemon-yellow, yellow, yellow-orange and brick-red (garnet,
russet-brown, burnt-orange, amber) also occur. The colour of the flower
(or, to be precise, the standard petal) is distinct and is cultivar-dependent.
The colour of the crescent portion is considered separately. As with any
colour characteristics, observation is quite sUbjective and can be difficult.
The use of a colour chart is recommended. Wings and keel are generally
yellow or orange-yellow in colour. In some accessions a darker orange or
purple blush could be found on wings.
(b) Standard crescent

The inside of the standard petal has distinct markings at the base from
which lines of different intensities radiate and this is usually referred to as
the crescent. The intensity of colour of the crescent ranges from complete
absence to a prominent pattern. Variation is also observed in the colour of
these markings between cultivars. Five gradations in the colour of the
crescent were reported, from the faintest (almost absent) to the most
prominent deep purple colour (Hayes, 1933; John et al., 1954). Usually
three expressions involving the standard crescent can be observed: pres-
ence or absence, compactness of the radiating lines and brightness of the
colour. The species belonging to the section Extranervosae have the
markings on the back of the standard. In species belonging to section
Ambinervosae, the markings appear on both surfaces of the flower. A.
batizocoi has a pink or purple blush on the back of the standard.
(c) Flowering
Groundnut is indeterminate in growth to a degree. Depending on geno-
type, environment and temperature, flowering starts at about 25 days after
emergence. As the daily mean temperature rises from 20°C to 30 °C, the
number of days required for the first flowering is reduced from 38 to 25 in
subsp. hypogaea and from 35 to 24 in subsp. fastigiata (Ono, 1979). The
most prolific flowering occurs between 5 and 11 weeks after planting,
depending on the duration of the cultivar and the season, with a high
degree of first-formed flowers producing mature fruits.
Usually four or five stages of flowering can be distinguished. Very few
flowers are produced at stage I, followed by rapid flowering in stage II. A
peak is reached at stage III, followed by a decline in the number of flowers
produced at stage IV. In some cases there can be two peaks of flowering.
Very few flowers that open after the first and/or second peaks produce
mature fruits, unless early flowers are prevented from functioning normally
due to some environmental stress (Bear and Bailey, 1973; Mohammad and
Khanna, 1932). Groundnut produces more flowers than the plant can
sustain to develop into pods. About 40% of the flowers fail to develop from
the outset, while another 40% produce only pegs (Smith, 1954). Less than
20% of the flowers produce mature fruit (Donovan, 1963) under best
conditions and sometimes less than 15% (Lim and Hamdan, 1984).
Genotypes which flower early show greater synchrony, and those which
produce most of the flowers during the first two weeks of the flowering
period produce greater numbers of pods (Sastry et al., 1985). Removal of
some flowers every day can prolong flowering (Bear and Bailey, 1973:
Seshadri, 1962).
On a given day, usually only one flower in an inflorescence reaches
anthesis, except in var. vulgaris in which two or more flowers may reach
anthesis on the same day. Intervals between successive flowers on an
inflorescence may vary from one to several days. A day before anthesis the
flower bud is 6-10 mm long. During the day the hypanthium elongates
slowly and the buds grow to 10-20 mm. The elongation is faster during the
night and at the time of anthesis the buds are about 50-70 mm long (Smith,
1950). Buds generally open at the beginning of the light period; it may be
delayed in cold or wet weather. The dehiscence of anthers takes place 7-8
hours before the flower opens or sometimes much later (Bolhuis et at.,
1965). Though the stigma was reported to be receptive from 24 hours
before to 12 hours after the opening of the flower (Hassan and Srivastava,
1966), generally it is receptive only a few hours before anthesis (Sastri and
Moss, 1982). On warm, sunny days most of the floral parts may wither
within 5-6 hours after flowering. Only the ovary and base of the style
remain turgid after the day of anthesis. Flowering generally occurs during
the early hours in the morning with some delay on cold/wet/dull days.
Generally, anther dehiscence occurs in the bud about one hour before
flowering. The pollen tube grows at the rate of about 1 cm per hour,
resulting in fertilization 5-6 hours after pollination (Lim and Gumpil,
1984).
3.5.8 Pollination and fertilization

Normally the stigma protrudes above the anthers and is receptive before
anthesis. As the stigma is enclosed in the keel and surrounded by the
staminal column, self-pollination is most common with a high frequency of
cleistogamy (Murty et al., 1980). Rarely, the stigma and anthers are
exerted from the keel and, as the stigma is receptive, some amount of
outcrossing may occur. Outcrossing ranged from less than 1.0% to 3.9%,
depending on the season, genotype and location, and several bee species
were found to be the pollen vectors (Culp et at., 1968; Gibbons and
Tattersfield, 1969; Hammons, 1963, 1964; Leuck and Hammons, 1969;
Srinivasalu and Chandrasekharan, 1958). Fertilization is complete in about
6 hours after pollination, and within 5-6 hours the flower may wither.
3.5.9 The peg

Generally fertilization is completed before midday. After fertilization, the
flower droops, the corolla closes, the calyx tube bends and the flower
withers. The ovary at the base of the calyx tube starts growing actively
within a week by the activation of the intercalary meristem present below
the ovary. The green ovary turns purplish from the tip downwards. The
developing ovary pierces through the floral parts to reveal an elongating
peg. The peg or carpophore is a stalk-like structure that carries the
fertilized ovules at its tip. Its growth is positively geotropic until it enters
the soil to some depth (up to 5-7 cm). The tip then becomes diageotropic
or almost horizontal. Only then does it begin to develop into a fruit. The
peg usually withers away if it fails to contact and penetrate the soil after it
extends to about 15 cm. In humid conditions, however, some var. fastigiata
cultivars can form aerial pods. These are underdeveloped, green and small.
True breeding aerial podding types have also been reported (Prasad, 1985)
and the physiology of such aerial podding, producing normal viable seeds,
needs further investigation.
The unique nature of the peg led to several misconceptions about its
homology and it has been described as a gynophore, a carpophore, or an
apetalous flower (Brennan, 1969; Jacobs, 1947). The peg has been defined
as the young fruit during the stalk-like phase of its development (Smith,
1950). The numerous minute plastids that develop after fertilization in the
epidermal walls of the peg were found to be responsible for its positive
geotropic movement (Patel and Narayana, 1935). The elongation of the
peg ceases as a result of auxin formed in its distal portion which moves with
strict basipetal polarity through the peg tissue and the intercalary meristem
(Jacobs, 1947, 1951a, 1951b). Sometimes, the peg or carpophore has been
incorrectly named gynophore (Smartt, 1976a). Bharathi and Murty (1980)
suggested that the peg is a highly reduced gynophore and is discernible as a
meristem at the base of the ovary. However, considering that the interca-
lary meristem basal to the ovary elongates and produces articulated pods in
most of the wild Arachis species, the peg is a carpophore (Ramanatha Rao,
1988).
(a) Colour
The colour of the above-ground portion of the pegs varies from shades of
pink to purple, in contrast to the normal green; and such colours are due
to anthocyanin pigments. Different peg colours such as light purple, dark
purple and green (no pigmentation) have been reported, as has a loose
association of the green colour of pegs with white testa colour (Varisai
Muhammad et at., 1973c). Though the development of pigment in the peg
is cultivar-dependent, it is greatly influenced by exposure to sunlight and
grades of pink or purple can occur on the same plant. Pegs can be broadly
classified as pigmented or green but the portion in the soil is generally
white (colourless).
(b) Hairiness
The developing pegs have minute white hairs that give a downy appearance
(Seshadri, 1962). These hairs are deciduous and are shed as pegs mature.
Above-ground parts of pegs may have stomata, lenticels and multicellular
trichomes whereas below-ground parts may bear unicellular structures
resembling root hairs which can reach very high density and may be up to
0.75 mm long. Similar structures may be found on the developing pod,
but at later stages these hairs degenerate and large lenticels may then
be present on pegs (Webb and Hansen, 1989). The species belonging
to section Extranervosae produce roots on the underground portion of
the peg.
(c) Size
The depth at which the pods are produced is determined by the extent of
the peg's penetration into the soil. Though significant cultivar differences
have been reported (Seshadri et al., 1955), variation within the same
cultivar for the size of the peg is also very great. The length of the peg was
reported to be closely related to the differences in branching pattern (Ono,
1979).
In wild Arachis species, the length of the peg is highly variable and can
be up to a metre in some species. In the context of distribution, the length
of the peg is important to produce the fruit as far away as possible from the
mother plant. A long peg, together with a long isthmus, can place the seeds
at a considerable distance from the mother plant. The orientation of the
peg in the wild species - horizontal or vertical - is an important taxonomic
criterion.
The thickness of the peg varies from about 1 mm to 2 mm. An assess-
ment of the thickness of the pegs in the different types showed that the
cultivars of subsp. fastigiata had thicker pegs than those of subsp. hypogaea
(Seshadri, 1962). The strength of the peg is an agronomic attribute govern-
ing the ease of harvesting of the crop.
3.5.10 The fruit

The fruit, referred to as a pod, is a slightly modified lomentiform indehis-
cent carpel. The cultivar differences in the proportion of pegs that pene-
trate the soil developing into pods were found to be significant (Seshadri,
1962; Singh et al., 1981). The mature pod normally contains up to four
seeds. Occasionally five or even six seeds per pod have been recorded.
Single-seeded pods may be produced when all the ovules except the
proximal abort (Smartt, 1976a). The pod size may range up to 8.0 cm x
2.7 cm. The fruit consists of valves, structurally dehiscent but functionally
indehiscent. Under pressure, pods split along the longitudinal suture. This
is the normal line of dehiscence and the mechanical tissue of the mesocarp
is interrupted along this line (Richter, 1899). The shell consists of an outer
spongy layer, a middle fibrous and woody layer, and an internal layer
which turns thin and papery upon maturation. Mechanical tissue on the dry
pod gives it a reticulate pattern with longitudinal ridges (up to 10) and less
prominent transverse ridges.
(a) Pod development

The ovule grows gradually up to flowering and the growth continues after
fertilization to some extent (Figure 3.6). The embryo, dormant during peg
elongation, begins to develop three to four days after commencement of
pod development (Murty et al., 1980). The basal ovule develops first. The
peg becomes diageotropic and almost horizontal when it reaches its
maximum depth in the soil and ceases to grow, and pod development
starts. The enlargement of the pod proceeds from base to apex. The pod
expands rapidly underground by the growth of a large parenchymatous
tissue (endocarp) lying between the ovules and shell layers (Schenk, 1961).
The pod grows to its maximum size in about three weeks after the pene-
tration of the peg into the soil. The endocarp recedes as the ovules grow
and disappears completely by the time the seeds mature. During this
period the inner face of the shell becomes increasingly darker in colour,
associated with an increased tannin content, and becomes very dark brown
on maturation (Gregory et al., 1973). It takes about 60 days from the time
of fertilization to full maturity (Patel et al., 1936). The size of the mature
pod and seeds is influenced by genotype, soil and method and time of
cultivation (Shibuya, 1935). The normal podding zone is located 4-7 cm
below the soil surface. The optimum soil temperature in the podding zone
is 31-33°C (Ono, 1979). Higher or lower temperatures prolong the dur-
ation of pod development. Lower soil temperatures (around 23°C) increase
the number of pods and pod weight but the filling period will be longer,
thus increasing the number of days to maturity (Dreyer et al., 1981).
(b) Number of seeds per pod

The number of seeds per pod may range from one to five or even six.
Single-seeded pods may occur in almost all cultivars but single-seededness
3
~
4
5
6
Figure 3.6 Pod development stages: 1. ovary after fertilization; 2. aerial peg, 5-7
days; 3. peg penetrating soil, 18-22 days; 4. pod enlargement, 14-21 days; 5. early
stage in pod development; 6. immature pod. (After Smith, 1950.)
is not genetic (Bunting, 1955) and it is usually a result of abortion of all but
the proximal ovules (Smartt, 1976a). Three-seeded pods are found in both
hypogaea and Jastigiata sUbspecies. The number of seeds per pod is
basically cultivar-specific, though it is influenced to some extent by season
and other factors (Seshadri, 1962). Three- or four-seeded pods may be
more common in bunch types than in semi-spreading and spreading culti-
vars, although two-seeded pods are the most common (Varisai Muhammad
et ai., 1973c). Cultivars belonging to var. Jastigiata possess predominantly
three- or four-seeded pods. The number of seeds per pod has been
extensively used for cultivar classification (Gibbons et ai., 1972; Varisai
Muhammad et ai , 1973c).
Most of the wild species, in contrast to the cultivated, produce articu-
lated pods. Generally they are biarticulate, though they may appear to be
uniarticulate since the isthmus between the articles quickly degenerates
with maturity of the proximal article.
(c) Beak
The tip of the indehiscent fruit may end in an appendage called the beak.
The prominence or presence or absence of the beak is dependent on the
cultivar and is a good diagnostic trait. Five grades can be used to classify
groundnuts based on this character: absent, slight. moderate, prominent
and very prominent. There can be some variation between the pods of the
same cultivar and the observation may be somewhat subjective; neverthe-
less, it is a very useful character. The shape of pod beak also varies but is
very difficult to define.
(d) Constriction
The constriction of the pod is an important character as it affects the
developing seed. Non-constricted pods generally have seeds with flattened
ends and are in contact with other seeds in the pod. Such a contact
sometimes results in embryo damage (Coffelt and Hammons, 1974). Seeds
from unconstricted pods tend to split during shelling and so cultivars with
no constriction are generally undesirable. On the other hand, pods with
deep constrictions tend to carry soil on them. They also break during
harvesting and shelling, reducing the market value. Most commercial
cultivars have some constriction between each seed; a few do not, espe-
cially the cultivars belonging to subsp. Jastigiata. Under abnormal con-
ditions (such as aerial fruiting), elongated pods with a filiform isthmus may
be produced by a cultivar that is normally only slightly constricted (Smartt,
1976a). In several wild species a filiform isthmus is characteristic, which is
developmentally complete, unlike the peg.
Pod constriction shows considerable variation both within and between
Anatomy 75
cultivars. However, the range of variation within a cultivar is not great
enough to prevent visual classification. Groundnut cultivars can easily be
graded into five groups based on the extent of pod constriction: none,
slight. moderate, deep and very deep.
(e) Reticulation
The mechanical tissue of the shell gives a reticulated or ridged appearance
to the pod. Usually the longitudinal ribbing is prominent but vertical ridges
may be prominent in some cultivars. Reticulation (venation, ribbing,
ridging) on the shell is an obvious visual characteristic. Typical var. fasti-
giata cultivars may be without any reticulation; however, some Peruvian
forms show very prominent ribbing. As with constriction, reticulation also
contributes to some extent to the cleanliness of pods at the time of harvest.
Five grades can be distinguished: smooth, slight, moderate, prominent and
very prominent.
(f) Size
Though the size of the pod is influenced to a some extent by soil and other
environmental conditions, it is an excellent diagnostic character. A range
of 8-20 mm in diameter has been reported and groundnuts have been
divided into five classes, depending on the mean length and mean pod
weight (John et ai., 1954; Varisai Muhammad et ai., 1973b). For a much
larger collection, a range of 11-83 mm for pod length and 9-27 mm for pod
diameter has been recorded (Ramanatha Rao, 1988). Ranges of 18-58 mm
for pod length, 9-19 mm and 7-39 mg for pod weight have also been
recorded (Retamal et ai., 1990). Pod length was found to be the most
useful diagnostic character in cultivar identification, in conjunction with
the other morphological characters (Salma, 1985).
In the case of the wild Arachis, the pod is usually articulated, the articles
being separated by the isthmus. The length of the isthmus is species-
dependant and varies from less than 1 cm to 30-35 cm. In the strict
botanical sense, the pod consists of the articles and the isthmus, but for
diagnostic purposes the size of a single article (preferably the apical) is
used.
3.6 ANATOMY
Groundnut yields are comparatively low, especially in the developing

countries. While the reasons for this are many, information on anatomy of
the various organs will help in understanding the physiological processes
governing higher yields. This information is likely to have direct bearing in
formulating research programmes in the disciplines of plant breeding,
agronomy, pathology, entomology and physiology. Efficient plant types
can be visualized in terms of disease, insect and drought stress, photosyn-
thetic efficiency and harvest index if more precise information on the
anatomy of various vegetative and reproductive organs is available.
Detailed accounts of groundnut anatomy can be obtained from many
works (Badami, 1935; Bharathi, 1981; Bharathi and Murty, 1984; Murty,
1988; Petit, 1895; Richter, 1899; Suryakumari, 1984; Suryakumari et at.,
1983,1984, 1989a, 1989b; Waldron, 1919; Yarbrough, 1949, 1957, 1957b).
3.6.1 Root
As groundnut roots typically lack an intact epidermis because of the lack of
specific epidermal initials, root hairs of the usual type rarely appear. Cells
originating in the lateral root cap region and in the outer cortex form the
surface layers of the root, the outermost of which is split and sloughs off.
Experiments have clearly established an active absorptive function of
solutes for this region (Richter, 1899; Yarbrough, 1949). Root hairs are not
entirely absent and are seen under special conditions and where humidity is
high and also more on lateral roots than on the primary root (Badami,
1935; Waldron, 1919, Yarbrough, 1949). The primary root has a tetrarch
vascular cylinder (Figure 3.7(c)). Meta and protoxylem tissues are differ-
entiated as in other dicotyledons. The large pith portion of the primary
root breaks down with age. During the early stages of seedling growth, the
cortex and pith contain abundant food stores. The root and hypocotyl
develop active cork and vascular cambium.
Lateral roots are formed by the activity of the endodermis and pericycle
which actively contribute cells to the new root apex (Petit, 1895; Richter,
1899). Lateral roots arise opposite the primary xylem plates. A four-
ranked arrangement of lateral roots is thus developed on the primary root.
The lateral roots are diarch and do not have a pith (Figure 3.7(d)). Thus
they differ from the primary roots of other dicotyledonous roots
(Yarbrough, 1949).
3.6.2 Stem
The description of the anatomy of Arachis stem can be found in several
earlier works (Reed, 1924; Waldron, 1919; Yarbrough, 1957b). The struc-
ture of the stem in general conforms to that of the typical dicotyledon type.
The epidermis of the stem consists of small cells with a thick cuticle. The
hypodermis is formed of angular collenchyma. The cortex consists of two
parts: the first part is chlorenchyma, and the inner part consists of larger
celled and thin-walled parenchyma (Yarbrough, 1957b). The vascular
Anatomy 77
Tetrarch stele
Figure 3.7 Structure of root and stem. Transections of ( a) primary stem; (b) stem
(secondary structure); (c) primary root; (d) lateral root with diarch stele and no
pith.
bundles are endarch and vary from 20 to 40. Five or six of them are larger
than the rest. Tannin cells occur in the stem (Figure 3.7(a)).
During the secondary growth of the stem, many pith cells break down
and form a cavity. An active fascicular and an interfascicular cambium
form considerable secondary tissue. Periderm develops in outer cortical
cells. Most cells of the cork cambium collapse before periderm develop-
ment (Figure 3.7(b)). Both the main axis and the cotyledonary laterals are
obconical in form when the primary tissues only are considered. The
number of fascicular bundles is greater in the upper internodes than in the
lower ones. The diameter of the pith is greater likewise (Yarbrough,
1957b).
3.6.3 Leaf
Routine information on leaf structure in Arachis was provided in earlier
studies (Reed, 1924; Yarbrough, 1957a). Studies on possible relationships
between leaf structure and resistance to leaf diseases were also made by
some workers (D'Cruz and Upadhyaya, 1961; Hemmingway, 1957;
Suryakumari et al., 1989a). A similar study was made by Pallas to find out
the reason for the abnormally higher conductance from the adaxial side of
the leaf than from the abaxial (Pallas, 1980). Detailed studies of leaf
structure in cultivated and wild species, interspecific hybrids and experi-
mental polyploids are also available (Suryakumari, 1984; Suryakumari et
al., 1983, 1989a).
(a) Cultivated varieties

Stomata
Stomata are present on both leaf surfaces with a higher frequency on the
adaxial surface, with few exceptions. Stomata are mostly of the paracytic
type but occasionally anisocytic, anomocytic and hemiparacytic types also
occur. Their frequency varies between 183 mm 2 to 302 mm 2 . Stomata vary
in length from 16-18 f.Lm (Suryakumari, 1984).
Epidermal cells
The epidermal cells are polygonal with straight anticlinal walls. In some
cultivars, the adaxial cells are larger than the abaxial cells.
Epidermal hairs
The epidermal hairs are non-glandular and uniseriate, with a thick basal
cell. The body of the hair consists of a short subterminal cell and a long
terminal cell. The hairs range in length from 0.02 to 0.92 mm. Epidermal
hairs occur on both surfaces of the leaves (Suryakumari, 1984).
Structure of lamina
Both the upper and lower epidermis are uniseriate with a thin cuticle. The
mesophyll is divisible into the spongy and palisade tissues. The palisade
cells occur beneath the upper epidermis only, loosely arranged in two
to four vertical layers. The poorly developed spongy parenchyma is
represented by a uniseriate layer. A layer of water-storage cells occurs
beneath the lower epidermis and the spongy layer. Tannin cells project
inwards from the cells of the upper epidermis. The thickness of the lamina
Anatomy 79
Bundle sheath extension

cells in T.S.
Spongy
parenchyma
(c) (d)
20li
Figure 3.8 Leaf transeetions: (a) subsp. Jastigiata; (b) subsp. hypogaea; (c) A.
hatizocoi; (d) A. pusilla.
varies from 100-250 f.Lm and the vertical extent of palisade tissue ranges
from 40-120 f.Lm (Suryakumari et al., 1983).
(b) Wild species

Wild Arachis species differ from the cultivated species in stomatal length,
frequency of hairs and magnitude of tissue ratio (i.e. the ratio of all other
tissues of the leaf to palisade height). A. batizocoi differs from the other
species in section Arachis in the number of stomata and hairs, length of
hairs, thickness of lamina, extent of palisade, palisade cell height and
palisade cell breadth. The two subspecies of Arachis hypogaea differ from
each other in the frequency of stomata, lamina thickness and palisade
parameters (Suryakumari etal., 1983) (Figure 3.8(a)-(d».
(c) Interspecific hybrids

Due to increased use of wild Arachis species in groundnut improvement,
especially for the incorporation of disease resistance, more and more stable
interspecific hybrids are becoming available. Interspecific hybrids at trip-
loid, tetraploid and hexaploid level show a gradual increase in leaf thick-
ness, palisade thickness and diameter with the increase in the ploidy level.
However, these characters deviate at the tetraploid level, the order being
4x > 3x < 5x < 6x.
(d) Polyploids
Diploid wild species of Arachis exhibit higher values than the tetraploid
species for the following parameters: frequency and length of hairs; fre-
quency of stomata; leaf thickness; and thickness and diameter of palisade.
In Arachis, features such as decreased size of epidermal cells, high fre-
quency of epidermal hairs, strongly developed palisade and higher percent-
age of water storage tissue were found to be associated with drought
resistance (Suryakumari, 1984).
3.6.4 Reproductive organs

Detailed accounts of microsporogenesis, megasporogenesis and repro-
duction in groundnut have been described by several workers (Badami,
1935; Banerji, 1938; Reed, 1924; Shibuya, 1935; Smith, 1956a, 1956b).
Similar studies of wild species have also been reported recently (Bharathi
and Murty, 1984; Suryakumari et at., 1989a).
(a) Cultivated varieties

For the most part, embryological features of A. hypogaea are of the
dicotyledonous type. In the following account, only the salient features are
given and the detailed routine descriptions are avoided.
Microsporogenesis and male gametophyte

The androecium consists of eight functional stamens and two sterile fila-
ments. In the functional stamens, four have oblong anthers, of which three
are tetrasporangiate and one bisporangiate (Figure 3.9(j)). The other four
stamens have bisporangiate globose anthers. A very young anther in
transverse section shows an epidermis enclosing a homogeneous mass of
cells (Figure 3.9 (a)-(c)). The development of anther wall corresponds.
to the dicotyledonous (basic) type (Davis, 1966; Xi, 1991). In a mature
anther, four wall layers - epidermis, endothecium, middle layer and tape-
tum - are distinct (Figure 3.9 (d)-(h)). Cells of the endothecium become
radially elongated and develop typical fibrous thickenings at the time of
meiosis in the pollen mother cells. The middle layer is ephemeral and
disappears during the maturation of pollen (Figure 3.9 (h)-(i)). In the
initial states, the tapetum is uniseriatewith uninucleate cells which become
bi- or tetranucleate with a vacuolated cytoplasm. Sometimes these nuclei
fuse to form polyploid nuclei (Figure 3.9 (k)-(n)). The tapetal cells are
glandular in nature. They degenerate before the dehiscence of the anthers
(Figure 3.9(0)).
The primary sporogenous cells divide mitotically to increase the number
of microspore mother cells which undergo normal meiosis. Occurrence
of cytomictic channels between microsporocytes, correlated with meiotic
Anatomy 81
C9~.~
~(b) ~~) (a) (c)
. 15!~. (k)
~()
.. (/)
i (n)
Figure 3.9 Microsporogenesis in Arachis hypogaea and male gametophyte (ar,

archesporial cell; e, epidermis; en, endothecium; gn, generative nucellus; ml,
middle layer; sp, sporogenous tissue; vg, vegetative nucellus): (a), (b) transverse
sections of anthers at successive stages of development; (c) a portion of young
anthers marked in (a) showing a homogeneous mass of parenchymatous cells; (d) a
portion of (b) enlarged to show hypodermal archesporial cells; (e), (f), (g) anthers
showing different stages of development of wall layers; (h), (i) mature anthers
showing meiosis in pollen mother cells, the disintegrating middle layer and fibrous
endothecium; U) one bisporangiate and three tetrasporangiate anthers; (k), (I),
(m), (n) tapetal cells showing one, two and four nuclei and also polyploid nucleus;
(0) another showing disintegrated middle layer and tapetum, and fully developed
fibrous bands in endothecium; (p) tricolpate 2-celled pollen grain. (After Murty,
1988.)
synchrony, has been reported (Xi, 1991). Cytokinesis is both successive
and simultaneous, divisions resulting in tetrahedral, T-shaped or isobila-
teral tetrads (Figure 3. 9(i) ). The microspores soon separate from each
other and each secretes its own wall, which differentiates into an outer thick
exine and inner thin intine.
The uninucleate microspore with centrally situated nucleus increases in
size. Due to the vacuolation of cytoplasm, the nucleus migrates to the
periphery and divides into a large vegetative cell and a small densely
cytoplasmic generative cell. The pollen grains are tricolpate and bicelled at
anthesis (Figure 3.9 (p).
Megasporogenesis and megagametophyte

The ovules are hemi-anatropous, crassinucellate and bitegmic. The micro-
pyle is formed by both the integuments. The archesporium in the ovule is
sub-hypodermal and is represented by a single cell. It directly functions as
the megaspore mother cell (MMC) (Figure 3.10 (a)-(c». The MMC
increases in size and divides meiotically to produce a linear, occasionally
T -shaped tetrad of megaspores (Figure 3.10 (d), (e). The chalazal mega-
spore undergoes three mitotic divisions and gives rise to an eight-nucleate
polygonum type embryo sac (Figure 3.10 (f)-(k». The mature embryo sac
is seven-celled with a three-celled egg apparatus formed of two synergids
and an egg cell. The synergids are pyriform in shape and have filiform
apparatus. The two polar nuclei in the central cell fuse to form a secondary
nucleus during fertilization. The antipodal cells at the chalazal end are
ephemeral and disintegrate by the time the embryo sac is organized (Figure
3.10 (k».
The nucellus disintegrates by the time the embryo sac reaches maturity.
However, some nucellar cells in the chalazal region become thick-walled
and form a hypostase, and at the micropylar end form an operculum. After
the disintegration of the nucellus, the cells of the inner integuments
adjacent to the embryo sac become radially elongated and densely cyto-
plasmic and form an integumentary tapetum or endothelium.
Fertilization
Fertilization in groundnut is porogamous. After pollination, one of the
synergids degenerates. The pollen tube enters the embryo sac through this
degenerating synergid soon after pollination (Murty and Rao, 1979).
Triple fusion precedes syngamy (Figure 3.11). Fertilization takes place
12-18 hours after anthesis.
Endosperm
Endosperm is ab initio nuclear. A fertilized embryo sac contains abundant
starch, which is a conspicuous feature during its early development. The
starch grains are mostly digested when the endosperm has 8-16 nuclei.
Free nuclear divisions in the endosperm continue until the embryo reaches
Anatomy 83
the early dicotyledonous stage. A higher number of endosperm nuclei
(about 5000) occurs at this stage (Figure 3.12 (a)-(f)).
Wall formation progresses centripetally. Cell wall formation in the
endosperm first takes place around the embryo and then gradually extends
(Figure 3.12 (g)). The entire endosperm does not become cellular and the
nuclei at the narrow chalazal end continue to lie embedded in a common
(a) (b)
(d) (e)
Figure 3.10 Megasporogenesis and mature embryo sac development in A. hypo-

gaea (ar, archesporial cell; dant, degenerating antipodal; mmc, megaspore mother
cell; sec.n, secondary nucleus; sy, synergid): (a) longitudinal section of an ovule
showing subhypodermal archesporial cell; (b), (c) enlarged megaspore mother cell;
(d) diad of megaspores; (e) megaspore tetrad; (f)-(j) successive stages of mega-
sporogenesis; (k) mature embryo sac.
cytoplasm. The coenocytic structure thus formed functions as an haustor-
ium. During the development of the embryo, the endosperm is absorbed
gradually and in the mature seed there is no endosperm.
Embryogeny
The embryogeny conforms to the Soland type of Johansen (Johansen,
1950). The first division of the zygote is transverse, resulting in an apical
cell and a basal cell (Figure 3.12 (c». These cells again divide transversely
(e) (I)
Figure 3.11 Fertilization in A. hypogaea (dsy, degenerated synergid; pen, primary

endosperm nucleus; pt, pollen tube; z, zygote): (a)-(f) successive stages in syngamy
and triple fusion.
Anatomy 85
cend
Figure 3.12 Endosperm and embryo development in A. hypogaea (cend, cellular

endosperm; ch, chalazal end; ned, nuclear endosperm; pen, primary endosperm
nucleus; z, zygote): (a) embryo sac in which syngamy and triple fusion are com-
pleted; (b )-( e) successive stages in the development of the embryo and endosperm;
(f) embryo sac showing globular embryo and endosperm with c. 3000 nuclei; (g)
embryo sac showing dicotyledonous embryo and cellular endosperm at micropylar
end.
and longitudinally and form a quadrant and then an octant embryo. At this
stage the embryo-forming region is clearly distinguishable from the suspen-
sor region by its smaller cells and denser cytoplasm. The embryo forming
region mostly divides by oblique walls and the basal cell, after two trans-
verse divisions, mostly divides by oblique walls and gives rise to a massive
suspensor. Localized divisions in apical meristems at two points at the
distal end of the globular embryo result in a bilobed heart-shaped embryo.
The later stages are marked by progressive increase in cells resulting in an
embryo with a massive radicle, two cotyledons and a massive hypocotyle-
donary region (Figure 3.12 (g». The embryo is straight and consists of a
robust radicle, a hypocotyledonary axis and an epicotyl made up of three
incipient shoot axes bearing nine or more differentiated leaves.
Seed coat
Initially the outer and inner integuments are two-layered. Periclinal
divisions take place in both the integuments, making them three-layered.
Further periclinal divisions occur in the derivative layers of the integument
only when the outer integument attains 11-13 layers. The outermost layer
of cells becomes columnar with thick walls, constituting the outer palisade
(Figure 3.13 (a)-(d».
Starch grains appear in the cells of the outer five to six layers. The cells
towards the inner integument do not contain starch grains and are com-
paratively small. During the later stages, the inner integument gradually
disintegrates (Figure 3.13 (e» and is completely obliterated in the mature
seed. Further divisions in the outer integument continue and in the mature
seed it is represented by 23-27 layers. The mature seed coat consists
of (1) a uniseriate outer palisade of columnar cells, (2) a middle layer of
parenchymatous cells, 20-25 cells thick containing starch grains and (3) a
single layer of isodiametric cells which represents the innermost layer of
the outer integument (Figure 3.13 (f» (Suryakumari, 1984).
(b) Wild species

Information on the reproductive behaviour of some of the wild species,
including diploids and tetraploids belonging to the section Arachis
CA. chacoense' Krap. et Greg. (nom. nud., PI 276235, 2n = 20); A. villosa
Benth. 2n = 20; 'A. duranensis' Krap. et Greg. (nom. nud., PI 219823,
2n = 20); and A. monticola Krap. et Rig. 2n = 40», the section
Rhizomatosae (A. hagenbeckii Harms. 2n = 40; and A. glabrata Benth.
2n = 40) and two undescribed tetraploid Arachis species (2n = 40), has
been provided in previous reports (Bharathi and Murty, 1984;
Suryakumari et at., 1989a).
No significant differences occur in the developmental morphology of the
species belonging to the different sections of the genus. This is true
especially of microsporogenesis and male gametophyte development and
of embryo sac development. However, minor differences occur. For
example, during megasporogenesis, the archesporial cell is hypodermal in
origin in the wild species. It divides into a primary parietal cell and a
primary sporogenous cell. The periclinal and anticlinal divisions of the
primary parietal cell produces a three-layered parietal tissue and the
primary sporogenous cell functions as megaspore mother cell (Figure 3.14
Anatomy 87
oi
(a)
(b)
(t)
(d)
Figure 3.13 Anatomy of seed coat in A. hypogaea (ii, inner integument; oi, outer
integument).
(a)-(h». Except in an undescribed tetraploid species, in which a triad was

observed, all other species have a linear or T-shaped megaspore tetrad.
Fertilization occurs more slowly in the wild species, since fusion of male
and female nuclei seems to be a slow process and the two nuclei are
detectable in close proximity to each other. The time taken for fertilization
varies from 6 to 48 hours. The number of free endosperm nuclei is much
less than in A. hypogaea at the dicotyledonous stage of the embryo.
In the seed-sterile species A. hagenbeckii Harms. the primary endo-
sperm nucleus divides only once. Thereafter the two daughter nuclei
(a) (b)
(c)
(d) (e) (f)
Figure 3.14 Megasporogenesis and megagametophyte in wild species (ar, arche-

sporium; end, endothelium): (a), (b) hypodermal archesporium; (c)-(h) successive
stages in the development of embryo sac; (i) degenerating fertilized embryo sac,
showing crushing of the embryo sac.
degenerate. No further development takes place. The embryo does not

develop beyond the four-celled stage: The endothelium becomes hyper-
plastic and crushes the embryo sac (Figure 3.14 (f)).
4.7 CONCLUDING REMARKS
The above morphological and anatomical descriptions of the groundnut

mainly consider the typical forms. However, several variants have been
References 89
reported from artificial or natural mutations or through hybridization
(Branch et at., 1982; Gopani and Vaishnani, 1970; Hammons, 1964;
Seshadri and Seshu, 1956). The morphological observations have been
mostly limited to the cultivated species. With the increased use of wild
species germplasm in studying and improving groundnut crops, there is a
need for extension of such description to wild Arachis and for the study of
the genus as a whole.
Large gaps still exist in our knowledge of the genus as a whole and more
investigations are necessary. Further anatomical studies, induding at least
a few samples of all 70 or so species, need to be carried out. These may not
only have taxonomical significance but may also help to identify any useful
association of morphological characters with anatomical features. Such a
comprehensive study of the genus as whole, which is presently lacking, will
be useful in gaining a better understanding of how wild species can be used
to improve the commercial groundnut crop.
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CHAPTER 4
Biosystematics and genetic resources

A.K. Singh and C.E. Simpson
4.1 INTRODUCTION
Biosystematics determines taxonomic status from experimental evidence

of the genetic diversity that arises within plant groups as a result of
evolution. It thereby assists applied biologists to utilize this diversity more
effectively. Without adequate analysis and assimilation of biosystematic
information, appropriate use of such diversity (by any method and in any
field) would be difficult and unpredictable. It is assumed that the evolution
which brings about differentiation between and within groups of organisms
has a built-in mechanism of isolation, the action of which is reflected in
sound systems of classification. Biosystematics has to uncover this through
appropriate investigations and provide unequivocal taxonomic nomen-
clature to the taxa which comprise the existing diversity. Given that
organisms each carry built-in information on the pattern in which their
development is ordered, it is the primary function of taxonomy and
biosystematics to utilize this information and produce systems of classifi-
cation and hypotheses to elucidate these patterns. The better such infor-
mation, the more useful it will be in answering biological questions,
particularly in relation to differentiation and the development of
divergence.
In Arachis, though interest in the use of genetic resources from both
cultivated Arachis hypogaea L. and wild Arachis species has increased and
extensive studies have been made of evolutionary relationships between
species of sect. Arachis and the cultivated groundnut (Smartt et ai., 1978a,
1978b; Singh, 1988; Singh et ai., 1991), taxonomic descriptions of all the
taxa of this genus are still not available, and confusion exists with regard to
the usage of botanical names and phylogenetic relationships among the
taxa (Resslar, 1980).
Groundnut genetic resources are available in the form of the naturally
evolved landraces of cultivated groundnut (A. hypogaea) in various centres
of diversity, breeding lines or material developed in different groundnut
Nomenclature 97
producing areas, and also the 70-80 wild Arachis species that are native in
South America. Together these present a satisfactory range of genetic
diversity. However, there may still be much more genetic variability in
centres of diversity that could be lost with the further expansion of
improved cultivars and the spread of modern civilization to remote areas
and peoples.
Significant progress has been made in characterization and evaluation of
groundnut germplasm. This has led to the identification of a number of
lines with desirable sources of resistance to both biotic and abiotic hazards
(Amin et at., 1985; Mehan, 1989; Subrahmanyam et at., 1990). But use of
the germplasm in genetic improvement of A. hypogaea is still in its infancy,
restricting progress in broadening the genetic base of the cultigen. In this
chapter we review the progress and present the current state of biosystema-
tics in the genus Arachis and of the available groundnut genetic resources,
highlighting promising areas for future research.
4.2 NOMENCLATURE
4.2.1 Subgeneric
The genus Arachis belongs to the Papilionoid legumes (Leguminosae,
Papilionoideae or Fabaceae), tribe A eschynomeneae , and sub tribe
Stylosanthinae. Based on morphological similarities, cross-compatibility
relationships and pollen fertility of interspecific hybrids (where available),
Krapovickas (1969, 1973), IBPGR (1990) and Gregory et al. (1973, 1980)
proposed a subgeneric classification dividing the genus into sections and
series (Table 4.1). However, none of these classifications were validly
published according to the rules of the International Code of Botanical
Nomenclature (Resslar, 1980). Therefore, all the sectional epithets which
have been proposed to date must be considered nomina nuda and have no
status under the code. Any of the proposed systems may be used equally
but the classification suggested by Gregory et al. (1980) has been used most
commonly, despite its non-validity, and has served well for defining broad
cross-compatibility relationships between taxa within the genus and
delimiting taxonomic boundaries. We follow the Gregory et al. (1980) plan
as modified by Krapovickas (1990) in the course of our discussion in this
chapter.
4.2.2 Specific and subspecific

The first Arachis species to be described was the groundnut itself
(A. hypogaea) , by Linnaeus (1753). Krapovickas and Rigoni (1960) and
Krapovickas (1969) divided this species into two subspecies, each of which
has two botanical varieties (Table 4.2). Gregory et al. (1980) suggested that
98 Biosystematics and genetic resources
TABLE 4.1 Taxonomic subdivisions of genus Arachis (Krapovickas, 1969, 1973,
1990; Gregory et aI., 1980) and proposed genomes
Sections Series Genome 2n=
Arachis 1. Annuae A,B,D 20

2. Perennes A 20
3. Amphiploides AB 40
Erectoides 1. Trifoliolatae E1 20
2. Tetrafoliolatae E2 20
Procumbensae P 20
Caulorhizae C 20
Rhizomatosae 1. Prorhizomatosae R 20
2. Eurhizomatosae 2R 40
Extranervosae Ex 20
Ambinervosae AM 20
Triseminalae T 20
the taxon, A. nambyquarae Hoehne, be considered only a variety of

subspecies hypogaea. In 1841 Bentham described some wild species includ-
ing A. villosa, A. tuberosa, A. glabrata and A. pusilla. Taxonomic revision
of the genus was attempted by Chevalier (1933, 1934, 1936), Hoehne
(1940) and Hermann (1954). Gregory and Gregory (1979) have estimated
that the genus consists of about 40-70 species. However, at present, only
23 species, one variety (A. nambyquarae) and a hybrid (A. batizogaea
Krap. et Fern., which was thought to have originated from a cross between
A. hypogaea and A. batizocoi) have been validly described and named
(Table 4.3). In addition, there are 12 invalidly named species which
regularly appear in the literature, such as A. cardenasii, A. duranensis and
A. chacoense (corrected spelling = chacoensis). According to Resslar
(1980), they should be designated by placing the abbreviation 'nom.nud.'
after the author citation. In the case A. villosa var. correntina (one of the
validly described varieties of A. villosa, which sometimes is also referred to
as A. correntina) biosystematic effort has solved the nomenclatural prob-
lem, showing that morphologically and also in cross-compatibility patterns
A. villosa and A. correntina are two distinct species.
A revision of the taxonomy of the genus is being prepared by
Krapovickas and Gregory to cope with the large number of new and
distinct accessions which have been collected in recent years. This mono-
graph should alleviate problems which have been compounded by recent
germplasm collections. The non-availability of published descriptions of
most of the Arachis species has resulted in assigning affinities wrongly,
especially by those not familiar with the genus, and others who continue to
add to the list of invalid names.
Systematics 99
TABLE 4.2 Classification of groundnut (Arachis hypogaea)
Subspecies Variety Botanical Branching Growth Seed/

type pattern habit pod
hypogaea hypogaea virginia alternate prostrate to 2-3

erect
hirsuta peruvian alternate prostrate 2-4
runner
fastigiata fastigiata valencia sequential erect 3-5
vulgaris spanish sequential erect 2
4.3 SYSTEMATICS
Systematic studies have been little appreciated by funding authorities, who

have been reluctant to support them because they were considered to be
basic taxonomic research and to have little practical value. These mislead-
ing perceptions are now changing because of the valuable contributions
such studies are making to wide hybridization in crop improvement. It has
become clear that crops like wheat and maize, which are well understood
biosystematically, have progressed more in their genetic improvement than
those crops which lack such information. Biosystematic information, par-
ticularly on phylogeny and cross-compatibility relationships between a
cultigen and its wild relatives, has greatly assisted breeders and cytogeneti-
cists involved in exploitation of interspecific variability for genetic im-
provement of cultivated species.
4.3.1 Classical
With the advent of new techniques our ability to understand the taxonomic
and phylogenetic relationships, even at the molecular level, has increased
and data are now available from several lines of investigation for integrated
analysis to produce a sound taxonomic system. However, a classical
approach, based on morphological similarities and dissimilarities, is still
primary and paramount. The genus Arachis is morphologically quite dis-
tinct from its close relatives because of its unique reproductive system
leading to production of geocarpic fruit. On the basis of similarities
in morphological features of leaves, leaflets, inflorescence and anthers,
Arachis has been placed in the tribe Aeschynomeneae and sub tribe
Stylosanthinae together with relatives such as Stylosanthes (Taubert, 1884).
The genus has been further divided into sections and series by Krapovickas
(1969,1973), Gregory et al. (1973, 1980) and Krapovickas (1990), based on
morphological features of the root system, orientation of peg, venation on
the standard petal and size of flowers. Cross-compatibility relationships
have been used to support many conclusions in this work (Table 4.1).
TABLE 4.3 List of validly described species of the genus Arachis (sub-generic classification after Gregory et a!., 1973; sectional
modification by Krapovickas, 1990)
Species Series Section

1. A. batizocoi Krap. et Greg. Annuae Krap et Greg. nom. nud. Arachis nom. nud
2. A. glandulifera Stalker
3. A. helodes Martius ex Krap. et Rig. Perennes Krap. et Greg. nom. nud Arachis nom. nud.
4. A. villosa Benth. "
5. A. villosa var. correntina Burkart
(A. correntina Krap. et Greg. nom. nud.)
6. A. diogoi Hoehne
7. A. hypogaea L. Amphiploides Krap. et Greg. nom. nud.
8. A. monticola Krap. et Rig "
9. A. batizogaea Krap. et Fern.
10. A. repens Handro Caulorhizae Krap. et Greg. nom. nud.
11. A. guaranitica Chod. et HassI. Trifoliolatae Krap. et Greg. nom. nud. Erectoides Krap. et Greg. nom. nud.
" N
12. A. tuberosa Benth.
13. A. benthamii Handro Tetrafoliolatae Krap. et Greg. nom. nud.
14. A. martii Handro "
15. A. paraguariensis Chod. et HassI.
16. A. rigonii Krap. et Greg. Procumbensae Krap. nom. nud.
17. A. marginata Gardner Extr~nervosae
Krap. et Greg. nom. nud.
18. A. lutescens Krap. et Rig.
19. A. prostrata Benth.
20. A. villosulicarpa Hoehne
21. A. burkartii Handro Prorhizomatosae Krap. et Greg. nom. Rhizomatosae Krap. et Greg. nom. nud.
nud.
22. A. glabrata Benth. Eurhizomatosae Krap et Greg. nom.
nud.
23. A. hagenbeckii Harms
24. A. angustifolia (Chod. et HassI.) Killip
25. A. pusilla Benth. Extranervosae Krap. et Greg. nom. nud.
Systematics 101
The description of 23 described species of the genus Arachis has been
considered by Gregory et al. (1980) as inadequate for developing a sound
morphological basis of species recognition. He pointed to limitations in the
herbarium specimens deposited in the major herbaria of the world, and to
the undue emphasis given to vegetative aerial plant parts which have strong
resemblances, even among taxa that are only distantly related; he said that
insufficient attention had been given to reproductive and subterranean
parts that are more stable, distinct and useful in differentiating valid
taxonomic entities. For example, the strong morphological resemblance,
particularly in leaf shape, between A. hagenbeckii, A. chacoensis and some
accessions of Erectoides and Arachis has led to erroneous identification of
one Erectoides accession as A. diogoi. Now that A. diogoi has been re-
collected, even the novice can correctly identify the mistakenly identified
Erectoides if provided with complete herbarium specimens for purposes of
comparison.
4.3.2 Biosystematics
Biosystematics is concerned with the evolution of isolating mechanisms
that have led to differentiation and genetic isolation of populations; hence
it is a valuable tool when combined with classical data in delimitation of
species and their recognition. It is easy to separate two taxa in the case of
complete isolation but this is more difficult and subjective when isolation
has been only partial. From an applied point of view, the information
based on the concept of biological species - i.e. that populations which
cross freely and are capable of free gene-exchange belong to a single
biological species and those which do not are outside it - is very important.
It defines quite satisfactorily the ease and extent of possible genetic intro-
gression and hence is of great applied value in strategic crop improvement
research. Because Arachis is predominantly self-pollinated, geographic
isolation and other extrinsic isolating mechanisms are expected to be more
important initially in differentiation and evolution of species than intrinsic
genetic isolation. The following biosystematic studies have been made to
promote understanding of the species and their differentiation.
(a) Cytotaxonomy
Ghimpu (1930) and Kawakami (1930) were the first to determine the
chromosome number (2n = 40) for A. hypogaea. Later, Husted (1933,
1936) identified a pair of small chromosomes and a pair of chromosomes
with secondary constriction and a satellite, as markers in the genomic
complement of A. hypogaea, designating them 'A' and 'B' respectively.
Smartt (1965), Smartt et al. (1978a, 1978b), Stalker and Dalmacio (1981),
and Singh and Moss (1982) observed many diploid species of sect. Arachis
with such an 'A' chromosome and A . batizocoi without such a pair. Smartt
et al. (1978a) first designated them as the A and B genomes respectively.
Singh and Moss (1982) confirmed statistically the distinction of two
genomes, and formed two clusters - one represented solely by A. batizocoi
and the other by the remaining taxa (of those analysed) of sect. Arachis.
The larger group was further subdivided into three sub-groups: A. cardena-
sii; A. duranensis and A. spegazzinii; and A. villosa and A. correntina
(Singh and Moss, 1982, 1984a). In taxa with 2n = 4x = 40, A. monticola
and A. hypogaea were found to be karyotypically very similar. Stalker
(1991) reported another species, A. glandulifera Stalker (A. spinaclava
Krap et Greg.), in sect. Arachis with a highly asymmetrical karyotype,
distinctly different from A and B genomes; he designated this as
the 'D' genome. Somatic complements of many more taxa from other
sections have been studied, although not in great detail, i.e. A. glabrata 2n
= 4x = 40 (Gregory, 1946), several other accessions of Rhizomatosae
(Singh, 1985b) and a number of accessions with 2n = 20 from sect.
Erectoides (Singh et. al., 1990). In these accessions, very little karyotypic
similarity with the species of sect. Arachis has been observed. In tetraploid
accessions of Rhizomatosae, several chromosomes were observed in
quadruplicate, indicating their auto tetraploid nature (Singh, 1985b). These
observations indicate that structural differentiation of chromosomes at the
diploid level and two independent courses of polyploidization - autopoly-
ploidization in section Rhizomatosae, and amphidiploidization in section
Arachis have been involved in differentiation and evolution of species in
the genus.
Meiotic studies in Arachis species have revealed that chromosome pair-
ing in most diploid species is regular, forming 10 bivalents (Smartt et al.,
1978a: Resslar and Gregory, 1979; Singh and Moss, 1982). Consistent
quadrivalent associations were observed only in tetraploid (2n = 4x = 40)
species of sect. Rhizomatosae, supporting their autotetraploid nature. Such
associations could also form as a result of fixation of a structural hetero-
zygote in rhizomatous clones of these accessions. A few quadrivalent
associations have also been observed in some accessions of A. hypogaea
(2n = 4x = 40).
(b) Cytogenetical
The degree of genetic and genomic isolation as a result of differentiation at
individual loci, through rearrangement of chromosome segments or poly-
ploidy (both auto- and allo-) , could result in cross-incompatibilities, either
partial or complete. Cytogenetical analysis of hybridization products, the
study of meiotic pairing, and pollen and pod fertility in hybrids may
provide real insight into the probable mode of speciation and evolution on
the one hand, and the possibilities of gene exchange for use by the
groundnut breeder on the other. The first viable interspecific hybrid
reported in the genus Arachis was between A. hypogaea (2n = 40) and A.
Systematics 103
villosa var. correntina (2n = 20) and was produced by Krapovickas and
Rigoni (1951). Since then, the cultivated A. hypogaea has been crossed
with as many as 37 species (Simpson, unpublished data) and crossing
between different species from both the same section and different sections
has resulted in production of more than 250 successful hybrids (Kumar et
al., 1957; Smartt and Gregory, 1967; Gregory and Gregory, 1979; Singh
and Moss, 1984a, 1984b; Singh, 1985b; Pompeu, 1977; Simpson, 1991a).
Such results have generally corroborated the cross-compatibility relation-
ships established by Gregory and Gregory (1979), who showed that,
despite ploidy differences, it is easier to produce intrasectional than inter-
sectional hybrids. They found that intrasectional hybrid pollen fertility at
the same ploidy level averaged around 30.2% in sect. Arachis; 0.2% in
sect. Extranervosae and 86.8% in sect. Caulorhizae. Intersectional hybrids
were completely female-sterile with very low or no pollen fertility; some
failed to flower at all.
Sect. Arachis, to which A. hypogaea belongs, contains a number of
diploid wild species reported to be resistant to several diseases of ground-
nut, such as rust, early and late leaf spots, and rootknot nematodes
(Abdou et al., 1974; Subrahmanyam et al., 1980, 1983; Nelson et al., 1989).
They are therefore important potential sources of genes for breeding for
disease resistance in A. hypogaea. Concerted biosystematic efforts have
been made to understand phylogenetic relationships among these species
of sect. Arachis (Smartt and Gregory, 1967; Stalker and Wynne, 1979;
Resslar and Gregory, 1979). Singh and Moss (1984a) performed a compre-
hensive genome analysis in sect. Arachis and reported that hybrids be-
tween the diploid species they studied, except those with A. batizocoi, had
near normal bivalent frequencies (9.1-9.8) with moderate to high pollen
fertility (60-91%). Hybrids between A. batizocoi and other species had
low bivalent frequency (5.2-6.9) and very low pollen fertility (3-7%).
These results confirmed genetic separation of A. batizocoi (B genome)
from the other set of species of sect. Arachis (A genome). Formation of
8-9 bivalents in some of the pollen mother cells (PMCs) of AB hybrids also
suggested that both genomes have a similar basic gene complement.
Chromosome pairing in triploid hybrids between A. hypogaea and the
diploid wild species corroborated the fact that the two genomes present in
A. hypogaea are common to the genomes found in diploid wild species
belonging to sect. Arachis. Furthermore, it indicated that A. batizocoi is
the closest diploid relative of A. hypogaea, probably more so to subspecies
fastigiata than to subspecies hypogaea, with homology to one set of the
genome (10 chromosomes) of A. hypogaea. The second set is homologous
to the other group of diploid species containing an A genome, forming an
average of 9 bivalents in their triploid hybrids involving A. hypogaea. The
tetraploid, A. monticola, crossed freely with A. hypogaea and showed
normal pairing of chromosomes in the hybrids, indicating that, biologic-
ally, A. monticola may just be the wild form of A. hypogaea.
In continuation of the above biosystematic studies, Singh (1986a, 1986b,
1988) further studied chromosome pairing, pollen fertility and pod fertility
in hybrids between cultivated tetraploid A. hypogaea (AA and BB) and
both synthetic autotetraploids and amphidiploids from AA and BB
genome diploid species of sect. Arachis. The objective was to verify the
above inferences and to identify the most probable ancestors of A. hypo-
gaea. Formation of higher mean bivalent associations in hybrids between
A. hypogaea X AA BB amphidiploids compared with A. hypogaea X AA
AA amphidiploids supported the earlier inferences that, most probably,
two sympatric related diploid species - one with AA and the other with a
BB genomic constitution - hybridized and produced A. hypogaea through
amphidiploidization. Arachis batizocoi and A. duranensis were considered
to be the most probable donors of B and A genomes respectively. There
was no complete pairing (i.e. 20 II) in these crosses to the amphidiploids,
since both diploid ancestors and the tetraploid cultivated species have
undergone a long process of evolution, leading to their further genetic
divergence. However, it is possible that the actual progenitor species of A.
hypogaea have yet to be collected, or may be extinct.
Very few intersectional hybrids have been produced between the diploid
species of sect. Arachis and sect. Erectoides, with and without embryo
rescue to overcome postzygotic incompatibility barriers (Gregory and
Gregory, 1979; Singh, 1989). However, a high level of sterility in these
hybrids has restricted the studies on chromosome pairing and pollen
fertility. Attempts to cross synthetic tetraploids of sect. Arachis and amphi-
diploid or natural tetraploids Of sect. Erectoides and sect. Rhizomatosae
respectively have virtually failed. Singh (1988) observed that crossing at
the tetraploid level was more difficult than at the diploid level. Gregory
and Gregory (personal communication) produced several hybrids between
diploid sect. Arachis annual types and complex amphidiploid hybrids
within sect. Erectoides. Stalker (1981) studied meiosis in hybrids between
two diploid (2n = 20) species, A. duranensis and A. stenocarpa (= A.
stenosperma) of sect. Arachis and an amphidiploid (A. rigonii x A. sp.
GKP 9841 = 4x) of sect. Erectoides and observed a high frequency of
bivalents indicating chromosomal homologies between the members of
sect. Arachis and sect. Erectoides. Stalker (1981) also analysed chromo-
somal homologies in some other complex hybrids produced by Gregory
and Gregory (1979) between sect. Erectoides and sect. Rhizomatosae
which suggested distant homologies between the chromosomes of the two
sections, and common ancestry.
The cytogenetical data in Arachis indicate that two chromosome series,
2n = 2x = 20 and 2n = 4x = 40, are present in the genus, though most
species are diploid with 2n = 2x = 20. Polyploidy probably arose twice in
the genus, independently, once in sect. Rhizomatosae through autotetra-
ploidization and again in sect. Arachis through amphidiploidization.
Cultivated tetraploid A. hypogaea (AA BB) could have originated via
Systematics 105
domestication of the wild tetraploid species A. monticota, which most
probably originated through amphidiploidization of an Fl hybrid between
a pair of species containing AA and BB genomes. The autotetraploid
genome of sect. Rhizomatosae has some homology with the A genome of
sect. Arachis and the E genome of sect. Erectoides, and so there are
substantially common chromosomes in all three of these sections.
(c) Chemotaxonomy
Seeligman and Krapovickas (Krapovickas 1973, Krapovickas et al. 1974)
were probably the first to look at the chemical variability in the genus by
chromatographic analysis of flavonoids in the leaves of Arachis species.
They detected more than 20 compounds, and chemical variation coincided
well with the centre of morphological variation of the genus (Mato
Grosso). There was no variation west of the meridian of 51 0 , which can be
considered as the axis of the area of distribution of the genus, as it runs
parallel to the Paraguay and Uruguay rivers. The presence of common
flavonoid chromatographic spots in A. hypogaea, A. monticola and diploid
wild species of sect. Arachis corroborated their close phylogenetic
relationship.
Several workers have studied both seed protein and isozyme profiles
through polyacrylamide gel electrophoresis (PAGE) and immunochemical
methods (Neucere and Cherry, 1975: Cherry, 1975; Klozova et al., 1983a,
1983b; Krishna and Mitra, 1988; Singh et at., 1991). Tombs and Lowe
(1967) noted that one of the major storage proteins, arachin, is polymor-
phic and they identified three forms of it. The gross fraction of protein
profile in the taxa of section Arachis did not differ much in overall mobility
patterns, and expressed considerable similarity even considering the differ-
ences in ploidy levels (Singh et al., 1991). This corroborates the inferences
of Klozova et al. (1983b), based on similarities of patterns resolved by
immunochemical methods (immunoelectrophoresis and double diffusion),
that protein patterns in Arachis have been relatively conservative in evol-
ution. Singh et at. (1991) have further indicated that the variation in
protein profiles of accessions of the same species is very low, particularly
in a small sampling of A. hypogaea cultivars. If this pattern holds true in
analyses of a broad spectrum of A. hypogaea, protein profiles may not be
of much practical value in differentiation and identification of cultivars, but
protein profiles can be of great value in differentiation and broad classifi-
cation of the genus and for species identification. These studies support
interspecific relationships consistent with the taxonomic scheme developed
by Krapovickas and Gregory (Gregory et al., 1980: Krapovickas, 1990),
particularly for the breakdown of the genus into sections. They also
support species relationships, and the probable ancestry of A. hypogaea
deduced from cytogenetical evidence (Singh, 1988).
Preliminary cytophotometric studies on amount of cDNA performed by
Resslar (1980) recorded 4.92-5.98 pg DNA per cell for diploid species and
nearly double that for tetraploids. Annual diploids have 1 pg less DNA
than perennial diploid species, and tetraploid taxa belonging to two A.
hypogaea subspecies and A. monticola show some small but consistent
differences. Restriction Fragment Length Polymorphism (RFLP) studies
carried out in the USA (Kochert et al., 1991) have shown only a very low
level of variation in a small sample of tetraploids that included A. hypogaea
cultivars from the USA and wild A. monticola. However, the wild diploid
species from sect. Arachis show considerable variation in their RFLP
bands, though the RFLP patterns of tetraploids were more complex than
those of diploids. Nevertheless, the two constituent genomes of amphidip-
loids could usually be distinguished. They suggested A. ipaensis, A. dura-
nensis and A. spegazzinii to be the most closely related diploid progenitors
of the tetraploid A. monticola and A. hypogaea, rather than A. batizocoi,
as indicated by cross-compatibility data cytogenetic studies and protein
profiles. This could be explained by assumption of certain structural
changes occurring at the microevolutionary level within the genomes of A.
hypogaea, and the genetic divergence these have caused in the molecular
sequences resolved by the probe and restriction enzyme combinations used
by Kochert et al. (1991). This is probably one of the limitations of RFLP
techniques, where early erroneous conclusions are possible, based on the
homology between fractions of genomes, before a broad array of material
has been analysed using all possible combinations. One needs to use
caution in drawing such conclusions on homologies before evaluating all
possible combinations of probe and restriction endonucleases to reveal
most of the genomic sequences. Critical investigations are still needed to
resolve such contradictions.
The studies above indicate that useful variation exists for chemical
characters in the genus. However, it is clear that the potential value may lie
in resolving the problems of classification between broad groups, which
may help in establishing species relationships and their delimitation, and in
assigning affinities to new taxa. There is obviously considerable scope for
genetic and molecular characterization of various genomes in the genus
using these biochemical techniques, and for using the information in
further classification and developing appropriate technologies for their
exploitation. Limited variation detected in A. hypogaea may be misleading
and the situation could well change following analysis of a broader spec-
trum of the available germplasm. Hence, the application of biochemical
techniques in groundnut is still to be developed to enable its utilization in
(1) resolving polymorphism among cultivars of groundnut, (2) fingerprint-
ing of cultivars and (3) identification of molecular markers associated with
desirable traits, which can be used in indirect screening of germplasm for
specific traits to aid in the production of improved new varieties.
From the above biosystematic studies, it is possible to establish tenta-
tively a series of genomes which parallel the Arachis sections. Some
Phylogeny 107
of the taxonomic entities are monotypic, such as sect. Triseminale with
Arachis sp. GKP 12881, and 12922 and series Prorhizomatosae of sect.
Rhizomatosae with A. burkartii, while sect. Caulorhizae is only rep-
resented by a pair of taxa. This provides a very narrow level of variability
for making inferences; nevertheless, the strong genetic barriers observed
between these taxonomic entities are sufficient to recognize the twelve
genomes shown in Table 4.1. These correspond to the section and series as
initially proposed by Gregory and Gregory (1979) and expanded by
Krapovickas (1990).
The sect. Erectoides comprises only diploid species which are classified
into two distinct series corresponding to two sub-genomes . In
Rhizomatosae, based on compatibility, it is proposed that tetraploid
Eurhizomatous species have two genomes, one with homologies to sect.
Erectoides and the other to sect. Arachis (Krapovickas, 1973; Gregory and
Gregory, 1979). However, the Rhizomatosae are assumed to be primitive
and it is difficult to see Arachis evolving from Rhizomatosae (if the section
is indeed primitive), as diploid rhizomatous A. burkartii is genetically and
physically isolated from other Arachis species. Singh (1985b), based on
karyomorphology (chromosomes in quadruplicate) and meiotic behaviour
(with quadrivalents), has indicated that tetraploid rhizomatous species
appear to be autotetraploids. Hence, it is more likely that genomic hom-
ologies reflected between sections Erectoides, Arachis and Rhizomatosae
on the basis of cross-compatibilities may be due to a pivotal genome
common to all three sections that have evolved in parallel, evolving sect.
Arachis and sect. Erectoides through genetic and chromosomal alteration
on the one side, and sect. Eurhizomatosae via polyploidy on the other.
This appears plausible because several annual species of sect. Arachis do
cross with the species of tetrafoliolate Erectoides (Gregory and Gregory,
1979; Singh, 1989) although only two crosses were successful without
embryo rescue techniques. In sect. Arachis, the genome analysis does not
conform with series classification. Series Annuae consists of species with
the B genome, sub-genomes of genome A, and another recently dis-
covered D genome, while series Perennes consists of only A genome
species. The series Amphiploides combines both A and B genomes. These
genomic classifications work best in the light of biosystematic evidence
collected to date and are subject to modification in case evidence from
further collection favours a different interpretation.
4.4 PHYLOGENY
Phylogenetically the genus Arachis has a number of characters in common

with Stylosanthes; more so than with any other related genus, although
some characters are not common to all species. Thus it is plausible to
suppose that Arachis evolved from a form such as Stylosanthes. Similarly
within the genus Arachis, based on morphological relationships, sect.
Erectoides, series Trifoliolatae with its trifoliolate leaves and Extranervosae
with its syncolpated pollen have been considered to be the most primitive.
A. guaranitica of series Trifoliolatae of Erectoides has basal flowers,
grouped in a cluster at the crown of the plant. They appear to have a
subterranean ovary, and a hypanthium 5 cm in length, which exposes the
androecium and stigma, protected by the keel petal, for pollination. All
Trifoliolatae and one species of the Extranervosae also have fruit produced
from the crown of the plant. In series Tetrafoliolatae all species have
flowers above the cotyledonary node (termed aerial for simplicity) in
addition to those with basal ovaries with proximity to the soil for fruiting
through decumbent branches. In other sections, such as Ambinervosae,
Caulorhizae, Rhizomatosae and Arachis, the flowers are normally aerial on
procumbent branches. But the trifoliolate characteristic itself may not be
primitive as the condition in Tetrafoliolatae is closest to that of Zornia in
the Aeschynomeneae. The peculiar fruiting habit may also not be primitive.
In addition, populations of A. tuberosa (a member of the Trifoliolatae)
have been collected with tetrafoliolate leaves; therefore the trifoliolate
form may be a simple genetic mutant, advanced and specialized in nature
and not primitive at all. However, the Extranervosae are clearly differen-
tiated from the rest of the genus by their unique pollen type, which is
supposed to be very primitive. Moreover, species of this section have
developed strong genetic barriers and only cross with species of sect.
Ambinervosae.
The fact that some species of the sect. Ambinervosae can be crossed with
some species of sect. Extranervosae and tetrafoliolate Erectoides can be
considered as indicative of affinity of the Ambinervosae between some of
the ancient sections and those more recently evolved.
Arachis, Rhizomatosae and Caulorhizae are the other derived sections.
The latter two have developed systems of vegetative reproduction. The
Rhizomatosae, which have tetraploid taxa, are thought to be the most
recent (Krapovickas, 1973). On account of crossing affinities, Krapovickas
(1973) and Gregory and Gregory (1979) presumed that some rhizomatous
diploids might have crossed with a diploid Erectoides followed by doubling
of the chromosome complement to give rise to the centrally located and
broadly distributed tetraploid (2n = 4x = 40) Eurhizomatous species. This
phylogenetic pattern is in agreement with the postulated evolution of
flowering and fructification types (Krapovickas, 1973). However, cytogen-
etic evidence does not fully support this hypothesis and to date no hybrids
have been produced between Prorhizomatosae and Erectoides. The loca-
tion of Prorhizomatosae and Erectoides is also problematic. The distance
between the closest collection sites of these two groups does not indicate
that they were close enough to hybridize by natural means to be credible
progenitors of Eurhizomatosae. Further biosystematic investigations are
needed to resolve phylogenetic relationships between these sections.
Genetic resources 109
The derived sections do not necessarily have a wide distribution. For
example, that of the section Extranervosae is only exceeded by sects.
Eurhizomatosae and Arachis in its geographic area. According to
Krapovickas (1973), it is reasonable to assume that the formation of aerial
flowers on prostrate branches is more recent than that of the flowers
grouped on the crown. The production of subterranean fruits and pegs
originating above ground is probably an ancestral character. The geo-
graphical distribution of these characters supports the contention that
prostrate plants have been the more effective in spreading to new areas.
The cultivated A. hypogaea, which is of primary importance, is phylo-
genetically very close to the tetraploid species A. monticoZa and to diploid
annual species such as A. duranensis, A. spegazzinii (a form of A. duranen-
sis) and A. batizocoi on the basis of morphological affinities, in conjunction
with cytological, genetical and biochemical affinities. They are also found
in the same region of northwest Argentina and southern Bolivia along with
primitive A. hypogaea (Valls et aZ., 1985) and have been considered an
immediate ancestor of A. hypogaea. Tetraploid A. monticola differs from
A. hypogaea in several characters, such as biarticulated fruits (a character
it shares with other species of the genus Arachis), a much smaller fruit size,
pegs as much as three times as long as those of A. hypogaea, very weak
peg-pod attachment and being able to survive without human inter-
vention.
4.5 GENETIC RESOURCES
The information that has been obtained by biosystematists in the genus

Arachis and in A. hypogaea is of considerable value for breeders involved
in improvement of the cultivated groundnut. Studies on cross-compatibility
are of particular importance as they have indicated the taxa that are most
likely to be accessible to the breeders for improvement of the cultigen.
Besides germplasm of A. hypogaea, the cross-compatible species of sect.
Arachis (which is the most recently evolved and evolving, section) provide
another reservoir of easily accessible genetic resources. Therefore the pool
of landraces and cultivars of cultivated groundnut which have evolved
under different levels of natural and human-induced selection pressure,
along with the wild species of sect. Arachis and other sections of the genus,
together provide a wealth of material for genetic modification and im-
provement of A. hypogaea.
The distribution pattern of wild species and the origin and spread of the
cultivated groundnut throughout South America and later to other
countries is not totally understood. The peculiar distribution patterns of
some taxa and the apparent wide gaps between some collections raise
many questions regarding probable evolutionary pathways. Incomplete
sampling of the total of potentially available material and the apparent
absence of important intermediate species and landraces (via extinction,
introgression or simply absence of collection) are factors that limit knowl-
edge of local evolution. The extremes of phenotypic variation within the
genus appear to be represented in our collections, but additional collec-
tions continue to expand the bulk of what is considered extreme, while new
collections from extreme areas of the range of the genus continue to make
our knowledge more comprehensive.
The genus extends over more than 2.6 million km 2 of the South
American continent, from north-east Brazil to north-west Argentina, and
from the south coast of Uruguay to north-west Mato Grosso south of the
Amazon; and from the base of the Andes to the Atlantic (Hammons,
1982). Many of the areas where additional taxa probably exist have not
been accessible. The extensive distribution of the genus, coupled with the
lack of adequate transportation networks and the harshness of the environ-
ment, make the task more difficult in the light of financial and political
considerations. A similar situation exists for collections from areas of early
introductions in Africa and Asia, which may still have useful genetic
diversity but which may be lost with further expansion of improved culti-
vars and modern technology into remote areas.
Significant progress has been made in the evaluation of genetic re-
sources, leading to identification of a number of lines from cultivated
A. hypogaea and wild Arachis species with desirable traits and sources of
resistance to both biotic and abiotic hazards that reduce groundnut pro-
duction worldwide (Amin et aI., 1985; Mehan, 1989; Subrahmanyam et al.,
1990). However, utilization of these genetic resources is restricted by lack
of information on reaction to other contemporary stresses, illustrating their
usefulness, and the detailed knowledge regarding the gene(s) mediating
them (it is essential to select appropriate breeding strategies for their
effective utilization).
4.6 GENE POOLS
The above sources of gen~tic diversity have been classified into four
gene pools (Wynne and Halward, 1989; Smartt, 1990). The first order
or primary gene pool consists of taxa belonging to the cultivated species;
the secondary gene pool consists of A. monticola. Arguments have
been presented that, based on crossability and fertility of hybrids
between them, A. monticola and A. hypogaea should be considered
as part of the same gene pool of a single biological species. Crossa-
bility studies have also indicated that sect. Procumbensae (which is essen-
tially sympatric with sect. Arachis throughout its distribution) produces
normal pegs and pods when cross pollinated with sect. Arachis but the
embryo aborts at an early stage of development, indicating evolution of
postzygotic barriers to hybridization during the course of evolution
Gene pools 111
Figure 4.1 Gene pools of genus Arachis.
(Singh, 1989). Therefore, in all probability Procumbensae has co-evolved

with series Perennes of sect. Arachis and must share genes with A.
hypogaea.
On the basis of these observations, the five or four gene pool theory fits
the genus Arachis very well (Figure 4.1). These pools may be described as
follows:
1. The primary gene pool consists of landraces of the cultivated groundnut
from the primary centre of origin and diversity in South America, and
cultivars and breeding lines developed in various groundnut-growing
countries around the globe. It may include A. monticola found in north-
west Argentina which has otherwise been considered as a secondary
gene pool.
2. The secondary/tertiary gene pool consists of diploid species that are
cross-compatible with A. hypogaea, despite ploidy differences, and are
placed in sect. Arachis with A. hypogaea and A. monticola.
3. The tertiary/fourth gene pool consists of sect. Procumbensae, which in
all probability co-evolved with series perennes of sect. Arachis and can
share genes with A. hypogaea on overcoming postzygotic barriers.
4. A fourth/fifth gene pool contains the rest of the Arachis species that are
cross-incompatible or weakly cross-compatible to sect. Arachis and are
classified into six other sections.
The chance of exploiting the third and fourth order gene pools will
depend upon advances in biotechnological techniques of genetic trans-
formation and somatic hybridization. The process will be time-consuming
and expensive. The most accessible gene pools will be the primary and
secondary ones and their actual breeding value is likely to be more
productive and somewhat more predictable. The characters of wild Arachis
species which have most immediate attraction to groundnut workers con-
cern resistance to diseases and pests, for which the variation within the
primary and secondary gene pools of A. hypogaea and A. monticola is very
limited.
4.7 CENTRES OF ORIGIN
The centre of origin of the genus Arachis was most probably in central
Brazil (Gregory et al., 1980). The geocarpic habit of the plant suggests that
long-distance dispersal has been along streams and rivers of the South
American continent. According to Gregory et al. (1973, 1980), the most
ancient species are found in higher elevations. Their immediate descendent
relatives occupy the next lower erosion surface and their distantly evolved
descendants occupy a still lower and more recent eroded surface. To
support this view, they found unique associations of botanical features with
highland conditions. Further, as seeds moved to lower elevations they
became isolated in major river valleys: thus different sections of the genus
evolved independently in a parallel fashion. Sect. Caulorhizae was isolated
and evolved in valleys of the lequitinhonha and Sao Francisco rivers,
Triseminalae in the Sao Francisco, Ambinervosae in the drainage system of
north-east Brazil, and series Prorhizomatosae in the Uruguay River valley.
Sect. Arachis probably evolved in the basin of the Paraguay or in the upper
reaches of the middle Amazon, which is invaded by the upper reaches of
the Paraguay. In both, the captured species are somewhat confined in the
uppermost part of the drainage systems, though they are widely distributed
in their original location. Species in the sect. Erectoides occur in the
extensive dissection of the Central Brazilian Planalto by both the Parana
and branches of the Paraguay rivers. Eurhizomatosae occur sympatrically
with Erectoides but extend beyond the Paraguay and Parana, south of
Centres of diversity 113
Corrientes, east into Sao Paulo and north of Rosario Oeste. However, the
belief that species in different sections are geographically isolated has
changed, since considerable overlaps in distribution between the members
of sections Arachis, Extranervosae and Rhizomatosae have been recorded
(Simpson, 1984; Valls et al., 1985) and major sectional groups of the genus
have been found with widespread distributions.
Cultivated A. hypogaea most probably originated in the region of
southern Bolivia and northern Argentina, since A. monticola, the only wild
tetraploid species that crosses with A. hypogaea, is found in this area.
Hence this is the region of the presumed centre of origin of cultivated
groundnuts (Krapovickas, 1969). Most of the diploid sect. Arachis species
that may have given rise to the segmental allotetraploid (A. monticolal
A. hypogaea) - such as A. batizocoi (Krapovickas et al., 1974; Smartt et al.,
1978a, 1978b; Singh and Moss, 1984a; Singh, 1986b, 1988), A. duranensis
(Seetharam et al., 1973; Gregory and Gregory, 1976; Singh, 1986b, 1988)
and A. spegazzinii (a form of A. duranensis; Kochert et al., 1991) - are also
distributed here. These observations support the conclusion that this re-
gion of southern Bolivia and northern Argentina was the centre of origin
and primary centre of diversity for A. hypogaea. This does not however
take into account recent archaeological evidence which could significantly
alter present perceptions.
4.8 CENTRES OF DIVERSITY
The genus Arachis is naturally restricted to Argentina, Bolivia, Brazil,

Paraguay and Uruguay in South America, although some species such as
A. repens, A. glabrata and A. pintoi are now cultivated elsewhere. Arachis
species are found from the mouth of the Amazon (0°) south across the Sao
Francisco and the Jequitinhonha, and into the mild temperate zone to 34° S
on the shores of the south Atlantic in southern Uruguay. Specimens have
been found west to the Parana River, south of Corrientes, and from the
Gran Chaco region of Argentina. The north-west limits of the genus
appear irregular but information may be incomplete because of lack of
accessibility to certain areas. The southern and eastern boundaries of the
multi-galleried Amazon Selva along a curved line south and south-
westwards from the mouth of the Amazon through Goias, Para, and
Central Mato Grosso to the north Beni in Bolivia form a tentative northern
boundary for Arachis species. From there it proceeds south along the
foothills of the Andes to Salta (Figure 4.2).
Arachis grows from sea level to 650 m above sea level on the Planalto,
from southern Mato Grosso to southern Goias, and to 1450 m a.s.l. near
Jujuy. It is found among vegetation types from broken forest to open
grassland. Its species grow submerged, among stones bathed with water, in
dry gravel, and in flood plain alluvium; they are found from semi-arid
locations to regions that receive an average of more than 2000 mm rain
annually, and in regions subject to great flood or intense droughts. They
grow most commonly in friable or somewhat sandy soil but also on soils
that vary from almost pure sand to reduced humic clays, sands of various
alluvia, and dark red humic latosols to pure lateritic caps and gravel.
Adaptation of wild Arachis species to such diverse conditions led to the
Erectoides
Extrallervosae
.-. Eurhizomatosae
---- Triseminalae
0000000 Prorhizomatosae
<>---0 Calilorhizae
Ambillerl'osae
- - - Procllmbel1sae
50° 40°
Figure 4.2 Geographical distribution of genus Arachis.

development of greater genetic variability and more combinations of
characters than those presented by the cultivated species. The above
conditions probably contributed to the development of geocarpy and
tuberiform roots of various forms, but the most important factor was
probably the harshness of the dry season. This and the need to escape dry-
season fires (an almost inevitable annual occurrence) contributed heavily
to tuberiform roots and geocarpy. Under a helping human hand, the
tuberiform roots were selected against in the cUltigen but the geocarpic
fruit was retained.
The highest number of species occurs in Brazil, where all eight sections are
found and to which country four sections of the genus Arachis are restricted.
Bolivia has the second highest number of species, followed by Paraguay,
Argentina and Uruguay. Most species occurring in Brazil are restricted to
the west central region but there is a group of endemic species in the semi-
arid north-east. Further differentiation in patterns of genetic variability in
different sections occurred as a result of their adoption of different ecological
niches where they were caught within a series of land uplifts during their
movements downstream in the associated drainage systems.
Sect. Arachis occurs from northern Goias and southern Para, south and
west to the base of the last-erosion surface exposed in the Pantanal
(100-175 m a.s.I.), along the Paraguay across north-central Bolivia to the
foothills of the Andes, where it was caught up in the Pleistocene uplifts and
where distinct new species mark the drainage systems, unlike the occur-
rence of entire sections that characterize the older Miocene system.
Genetic isolation is not as strongly marked between most species of sect.
Arachis compared with those of other sections.
The probable region of origin of the cultivated groundnut (discussed
above) presents a wide range of distinct environments and there is evi-
dence that natural hybridization and the establishment of recombinants
have occurred in these areas.
Archaeological evidence suggests that groundnut has been cultivated for
over 3500 years. It was probably first domesticated in northern Argentina
and eastern Bolivia. It was subsequently introduced to Africa, India and
the Far East by the Portuguese; and from the west coast of South America
to the western Pacific to Indonesia and China by the Spaniards in the early
sixteenth century. It is possible that the ground nut may have travelled to
China well before the time of Columbus but additional evidence is needed
to support this idea. There were introductions to East Africa from Asian
countries. By the middle of the sixteenth century the groundnut was
introduced to North America and to other parts of the world. Since this
period of introduction numerous forms have evolved (Hammons, 1982).
The introductions from Bolivia to Africa and Asia are more recent.
In South America, where the greatest amount of genetic diversity is
found, Krapovickas (1969) and Gregory and Gregory (1976) recognized six
gene centres for cultivated ground nuts (Figure 4.3). These centres are:
1. the Guarani region
2. Goias and Minas Gerais (Brazil)
3. Rondonia and north-west Mato Grosso (Brazil)
4. The eastern foothills of the Andes in Bolivia
5. Peru
6. North-eastern Brazil.
It is now apparent that at least one more primitive centre exists in

Ecuador, where a distinct group of landraces is grown at present.
These centres of diversity contain a high level of genetic variation due to
introgressive hybridization followed by human selection, resulting in pro-
duction of typical hybrid swarms, which might have been produced by
crossing between types with divergent characters. However, most can still
be linked with specific subspecies and botanical varieties. The Guarani
region is dominated by erect valencia types (subsp. fastigiata); subsp.
hypogaea is very rare here. There is little evidence of introgression be-
tween the two subspecies in this region, but hybrid swarms of intermediate
races between two botanical varieties of subsp. fastigiata do exist, and both
var. fastigiata valencia, Porto Alegre, and var. vulgaris Negrito, spanish,
are identified. It is possible that the spread of valencia types to other parts
of the world has occurred from Paraguay or central Brazil but a more likely
point of embarkation is from the north-east coast (i.e. the Amazon). The
Guarani region is also the centre of variation for the variety vulgaris, and
spanish types were probably also disseminated from this region (Krapo-
vickas, 1969; Gregory and Gregory, 1976).
The second region of Goias and Minas Gerais has a varietal pattern
distinct from the Guarani, but still is dominated by erect groundnuts
(subsp. fastigiata) , with very few examples of subsp. hypogaea. Cultivated
races belong to both varieties, fastigiata and vulgaris, without much indi-
cation of introgression. Rondonia, the third region of diversity, is typically
represented by the nambyquarae type of subsp. hypogaea and the Mato
Grosso by a yellow testa, erect type and A. villosulicarpa. The fourth
region in the eastern foothills of the Andes is a great centre of variability of
subsp. hypogaea var. hypogaea with few races of var. fastigiata. In Bolivia
there are indications of introgression between the two subspecies, and
'Overo' and 'Cruceno' types of groundnut are probably the product of such
introgression (Krapovickas, 1969; Gregory and Gregory, 1976).
From Peru, three distinct types of groundnut are collected, one of which
is like that found in pre-Columbian tombs: it has fruit with prominent
constrictions, veins and beak and belongs to subsp. hypogaea. This is called
the 'chinese' type in the USA. A second type, with similar fruit characters
but belonging to subsp. fastigiata var. fastigiata, has also been found in this
region. These two together were called the 'peruvian' type by Dubard
(1906). A third type belongs to subsp. fastigiata var. fastigiata but has
smooth pods with three to five seeds and almost no beak.
Distribution of wild Arachis species

Primary centre or origin of A.
hypogaea
o Secondary and tertiary centre of

diversity
Figure 4.3 Centres of origin and diversity of Arachis hypogaea in South America
(adapted from Gregory and Gregory, 1976). I A. hypogaea fastigiata var. fastigiata
and var. vulgaris; II A. hypogaea fastigiata var. fastigiata; III A. hypogaea hypogaea
var. hypogaea; IV A. hypogaea hypogaea var. hypogaea, A. hypogaea fastigiata
var. fastigiata; V A. hypogaea hypogaea var. hypogaea and var. hirsuta, A. hypo-
gaeafastigiata var. fastigiata; VI tertiary A. hypogaeafastigiata and var. vulgaris.
Williams (1991) studied the region of the north Beni of Bolivia/Peru. He
collected some extraordinary types which appear to be intermediate be-
tween subsp. hypogaea and subsp. fastigiata.
The sixth centre, north-east Brazil, is regarded as a tertiary centre with
all types. The Ecuadorian type is similar to the var. fastigiata types of Peru
but morphologically distinct and might even be considered intermediate
between vars. fastigiata and hypogaea.
Africa has been regarded as another important tertiary centre of diver-
sity. Introductions from Brazil probably became established concomitantly
with the slave trade in West Africa, while the east coast probably received
material which had come from the west coast of South America via the
Philippines, China and India. More recently Smartt (1990) suggested
significant introductions from Bolivia, India and China can also be con-
sidered as sources of much variability for cultivated groundnut at various
times.
4.9 COLLECTIONS
Collection and assembly of groundnut genetic resources has been exten-

sive. It received much impetus from the establishment of the International
Board for Plant Genetic Resources (IBPGR) in 1974, and the inclusion of
groundnut in the mandate of the International Crops Research Institute for
the Semi-Arid Tropics (ICRISAT) in 1976. Before this, extensive ground-
nut collections were maintained only at the Southern Regional Plant
Introduction Station, USA, at Agricultural Research Stations of the USA,
and in countries including Brazil, Argentina, Senegal, Israel, Taiwan,
India, Nigeria, Malawi and Zimbabwe.
With the establishment of the Genetic Resources Unit at ICRISAT,
various collections could be assembled at a central place for use by the
international community. The world collection was also enhanced through
specific collection expeditions to various groundnut-growing regions of the
world and to centres of diversity in South America (Figure 4.4). In these
expeditions various international agencies, such as IBPGR, and national
programmes, such as those of countries of exploration, and the United
States Department of Agriculture (USDA) have collaborated very closely
(Simpson, 1982, 1984), leading to the establishment of the world's largest
repository of groundnut germplasm at ICRISAT and in the USA under the
USDA. ICRISAT and the USDA now maintain around 14000 and 7545
accessions respectively, and provide basic genetic stocks to the inter-
national scientific community for further improvement of the groundnut.
They include both cultivated and wild Arachis species. The most complete,
accessible catalogues of collections are also the ones maintained at USDA
and ICRISAT with information on 7545 and 12 160 accessions respectively.
Many of the collections held at different places are duplicated through
Collections 119
exchange of material, and the total number of different genetic types is less
than that represented by the above numbers.
Systematic collection expeditions in South America were started by
Archer in 1936 (Valls et al., 1985). Besides the collection of landraces of
cultivated groundnut, efforts were made in these expeditions to collect
distinct Arachis species (Simpson, 1982). The expedition undertaken in
1959 resulted in the collection of six old and five new Arachis species. From
1961 to 1967, five new and three old species were collected. A species
belonging to sect. Ambinervosae was collected for the first time in 1967. A
1968 expedition resulted in collection of one old, one new and several
undescribed and unnamed species from sect. Erectoides. Many more new
wild Arachis species were collected in expeditions from 1976 to 1979; most
of these were from Bolivia and the edge of the Pantanal in Brazil. The
1976 expedition collected typical A. diogoi and re-collected A. lutescens. In
1980-81 a species referred to in some literature as A. sylvestris was
collected for the first time. New Ambinervosae species were discovered in
Brazil along with several new species of sect. Arachis and sect. Erectoides
(TriJoliolatae), and sect. Procumbensae in west and central Brazil. The re-
collection of A. tuberosa, A. lutescens and A. prostrata occurred in 1982.
A. marginata was re-collected with additional unnamed species of sect.
Extranervosae and annual species of sect. Arachis penetrating into north-
east Brazil, further extending our knowledge of the are.a of their natural
occurrence. In 1983, two new species from sect. Extranervosae and sect.
Erectoides (TetraJoliolatae) were collected. Each expedition provided addi-
tional recorded locations of both wild and cultivated groundnut, as a result
of which the map of natural occurrence for the different sections waS
greatly modified from the earlier one presented by Gregory et al. (1980)
(see Simpson, 1984). Figure 4.2 illustrates the current perception of the
distribution of Arachis species. The ranges of several sections, such as
Erectoides and Ambinervosae, were expanded due to widespread distri-
bution of A. sylvestris from the Ambinervosae, and due to discovery of a
TetraJoliolatae species east of the previously known range and of new
species of sect. Procumbensae in east Bolivia and north of the Pantanal
(Simpson, 1984).
The area of sects. Ambinervosae and Triseminalae also showed signifi-
cant expansion south to the valley of the Jequitinhonha, overlapping with
Caulorhizae and Extranervosae. The monotypic series Prorhizomatosae
was also extended to the east coast of Brazil.
Table 4.4 lists the number of accessions available at some of the major
centres. The collections are non-discriminatory in the sense that both
landraces and materials developed by breeders and/or released cultivars
were collected, since any present-day genotype could contain genes that
may be of use in the future. These collections can be classified into:
Figure 4.4 Groundnut-growing regions in the world.
Collections 121
TABLE 4.4 Status of Arachis germplasm at different gene banks
Resource ICRISAT Griffin Texas NCSU Campinas b

Georgia A&M
Accessions 7817 7545 3403

Landraces 5146 - a
Named cultivars 230 75
Breeding lines 4553 11498
Genetic stocks 131
Wild Arachis spp. 195 498 798 294 596
Taxonomic representation:
Section
Arachis 12044 15204 147 137
(wild)
Erectoides 21 68 32 68
Caulorhizae 2 34 2 60
Rhizomatosae 90 208 76 71
Extranervosae 2 63 5 178
Triseminalae 1 7 2 7
Procumbensae 0 30 29
Ambinervosae 0 27 2 46
Interspecific
derivatives 165 261 705
Unknown 1651 28
a Information not available

b Instituto Agronomico. Campinas, Brazil.
• Accessions: world collection assembled from other agencies with an

accession number.
• Landraces: non-uniform material (unselected) collected from farmers'
fields or purchased in local markets.
• Breeding line: material developed by breeders but not released as
cultivars.
• Genetic stocks: genotype identified by special features or sources of
resistance to biotic and/or abiotic stresses.
• Wild species: Arachis species belonging to one of the eight sections of
genus Arachis.
The assembly of such large collections is an excellent example of co-
operation between various international, national and state agencies.
However, there are several gaps in these collections of both wild and
cultivated species, as mentioned previously. According to earlier priorities,
major areas have been explored, and now countries and regions with
limited groundnut cultivation but which may still hold extensive variability
TABLE 4.5 Priorities for future collections
Country Areas to be covered Justification
Argentina 1. North and north-west To collect A. monticola

2. Corrientes and Entre Rios To collect A. villosa
Brazil 1. Rios Tapaj6s and Xingu Unexplored
2. North of Ilha do Bananal Unexplored
3. The valley of Rio Parnaiba To collect sect. Arachis and
A. hypogaea
4. Rio Parana to coast of Brazil To collect sect. Arachis and
and Rio de Janeiro to A. hypogaea
Uruguay
5. City of Campina Grande To collect A. martii
6. Northern part of Mato To collect A. villosulicarpa
Grasso and A. hypogaea
7. North-west part of Goias, To collect additional
Maranhao, northern Piauf germplasm of Arachis and
state A. hypogaea
8. Ceara, Rio Grande do Norte Landraees of A. hypogaea
and Paraiba states
9. South-eastern Amazon
region
10. Rondonia and Acre States
Bolivia 1. Northeast Bolivia along the To collect sect. Arachis and
valleys of Mamore and Procumbensae
Guapore
2. Bafiados del Izozog
3. Eastern Bolivia
Paraguay 1. East of Paraguay river To collect lost species
2. West of Paraguay river and To collect sect. Arachis
Rio Paraguay valley
3. North Chaco of Paraguay To collect sect. Arachis
Uruguay Along Uruguay river To collect A. villosa and
landraees of A. hypogaea
should be explored to make the collection more comprehensive. It is

important to do this before germplasm is lost due to drastic changes in
agricultural patterns. This is especially true for South America, which has
both wild and cultivated groundnut and where natural habitats are increas-
ingly being disturbed. Table 4.5 presents priorities for future collections,
partially based on an IBPGR paper by Simpson (1990).
4.10 GENETIC VARIABILITY
The wide range of genetic diversity in the cultivated and wild Arachis
species contradicts the earlier apprehensions of lack of variability.
Genetic variability 123
TABLE 4.5 cant.
Country Areas to be covered Justification
Peru l. South and southwest To collect jastigiata type

2. Central and east To collect jastigiata type
3. North and north-east To collect jastigiata and
hirsuta types
Ecuador Western half To collect Zaruma types
Colombia West and north Unexplored and
unrepresented
Mexico Whole country Landraces and hirsuta types
Caribbean Whole countries Unexplored
islands
China Unexplored Landraces and hirsuta type
Kampuchea Whole country Unexplored and
unrepresen ted
Laos Whole country Unexplored and
unrepresented
Thailand Wholc country Undcrexplored
Vietnam 1. North and northeast Landraces and hirsuta typc
2. Southern state Landraces
Bangladesh Whole country Unexplored and
unrepresen ted
Nepal Interior regions Landraces
Central African Whole country Landraccs for humid tropics
Republic
Gabon Whole country Early introduction zone
Angola Whole country
Republic Whole country Isolated
Malagasay
Cameroon Whole country To re-collect lost landraces
Namibia Coastal regions Early introduction zone
Senegal Whole country To re-collcct and preserve
lost variation
Together they present a considerable amount of diversity. At ICRISAT,

and at most other major repositories, a multidisciplinary approach is
adopted for characterization of groundnut germplasm, conducted by a
team of scientists with expertise in different disciplines who follow a
common groundnut descriptor list developed by IBPGR/ICRISAT (1981)
and revised subsequently. For these characterizations, emphasis has been
given to characters considered to be highly heritable. The different mor-
phoagronomic characters recorded are listed in Table 4.6. These descrip-
tors have also been applied to the 2000 most recent collections from South
America (Simpson et al. 1992). Other centres have applied these and other
TABLE 4.6 List of characters recorded
I Morphological characterization
1. Life form
2. Growth habit
3. Branching pattern
4. Stem pigmentation
5. Stem hairiness
6. Peg colour
7. Standard petal colour
8. Standard petal markings colour
9. Leaf colour
10. Leaflet length
11. Leaflet width
12. Leaflet shape
13. Leaflet hairiness
14. Number of seeds per pod
15. Pod beak
16. Pod constriction
17. Pod reticulation
18. Pod length
19. Pod width
20. Seed colour, primary
21. Seed colour, secondary
22. Seed colour, type of variegation
23. Seed length
24. Seed width
25. Seed mass
II Preliminary evaluation
26. Days to emergence
27. Shelling percentage
28. Days to maturity
III Further evaluation
29. Oil content
30. Protein content
31. Reaction to abiotic stresses.
32. Reaction to biotic stresses.
descriptor lists to large numbers of germplasm accessions. For example,

Pietrarelli et al. (1985a, 1985b) evaluated all the germ plasm ever collected
in Peru and Bolivia; and Howard et al. (1985) applied IBPGRlICRISAT
descriptors to a collection of South American germplasm lines. Further
evaluation of germplasm for additional descriptors such as reaction to
diseases, insect pests, drought and nutritional components has also been
carried out at ICRISAT and other locations.
The available germplasm (at ICRISAT as well as other centres) has
exhibited a wide range of vanahon for morphoagronomic characters.
Growth habit in wild Arachis species and the cultigen have striking simi-
larities: a relatively short main axis and four or more horizontal branches
arising from the axil of the cotyledon and the first pair of foliage leaves.
Further branching is basically of two types - alternate and sequential. In
the alternate type, pairs of axillary vegetative branches are produced
alternating with one or two axillary inflorescences; in the sequential type,
either a distal sequence of inflorescences or an unbroken sequence of
axillary inflorescences is present. Various combinations of the two also
occur frequently. A. monticola has both types (Gibbons, 1966) and so
has A. hypogaea. As much as one third of the germplasm collection of
A. hypogaea is a mixture of alternate and sequential, either within or
between plants.
In growth habit, cultigens can be of the procumbent runner type with
short (generally) or long main axis and laterals growing horizontally to
various lengths. Other genotypes may have decumbent laterals which have
an ascending tendency or are erect with shortened internodes in both
alternate and sequential branching patterns. These have been designated
as D1, D2, D3 and E, according to the angle between main axis and
branches. Spanish and valencia are mostly erect while virginias are gener-
ally spreading. Valencia and spanish genotypes have few secondary and
tertiary branches, while virginias have many such branches. A consistent
difference between the two subspecies and botanical types is that spanish
and valencia types have inflorescences on the main axis, while pure
virginias generally do not. However, recent extensive hybridization has
resulted in several intermediate types.
Leaves of virginia types are generally darker green than those of spanish
or valencia. In diploid species also, the annuals have lighter green leaves
than the perennials, though there is great variation in the depth of colour.
Mutations, recombinations and their segregants have evolved many more
types of leaf shape and colours.
The range of flower colour in Arachis is not great. Flowers are usually
orange or yellow, with or without marking, but certain accessions have
white and even brick red flowers (the latter only in A. hypogaea). Pods in
wild Arachis species are mostly lomentiform. The isthmus between the
seeds can vary from a few millimetres to greater than 10 cm (Gregory et al.,
1980). The cultigens have great variability for pod characteristics due to
different selection pressures for this commercially important trait.
Genotypes differ from deep to almost no pod constrictions, and from
prominently reticulated to essentially smooth pod surfaces. Some have a
distinct beak while others have no beak at all, and pod length varies from 1
to 9 cm. Similarly, there is considerable variation in seed size. Wild Arachis
species, A. cardenasii and A. correntina, have very small seed (but by no
means the smallest), weighing about 20-25 mg each. Some of the smallest
seeded genotypes of A. hypogaea may approach this level, while the largest
seed may weigh as much as 1.3 g each. Cultigens also show great variability
for testa colour, varying from white to tan, purple, dark red and black,
with different types of variegation or self colours.
This range of variability recorded for 12 160 accessions at ICRISAT is
summarized in Table 4.7. Further screening of groundnut germplasm to
assess the potential - particularly their reaction to different biotic and
abiotic stresses - has been carried out worldwide, resulting in identification
of a large number of accessions with desirable traits and sources of resist-
ance to many stress factors. The results of such screening at ICRISAT and
other major centres are summarized in Table 4.8.
Wild Arachis species have shown great variability in both perennial and
annual life forms, sequential or alternate branching patterns, and open or
compact prostrate habit. The reaction of wild species to various groundnut pests
and diseases has shown greater variation than is present in cultivated groundnut
and they provide a large number of sources for introgression of disease
resistance. Species such as A. glabrata, A. hagenbeckii, A. repens (Gibbons and
Bailey, 1967), A. chacoensis, A. villosulicarpa,A. sp. GKP 10596 (Abdouetal.,
1974; Fosteretal., 1981; Kolawale, 1976) and A. stenocarpa (= A. stenosperma)
(Sharief et al., 1978) were reported to be resistant to early leaf spot. A.
cardenasii and several other taxa from section Arachis (Kolawale, 1976),
Caulorhizae, Ambinervosae and Rhizomatosae (Abdou et al., 1974;
Subrahmanyam et al., 1985b) were found resistant to late leaf spot. Many
accessions of Arachis species have been found resistant to rust.
Many wild Arachis species have been identified as resistant to viruses. A.
glabrata and A. repens were reported to be symptomless carriers of ground-
nut rosette virus (Gibbons, 1969). A. pusilla (GKP 12922 probably not
authentic A. pusilla, better designated Arachis sp.), A. correntina (GKP
9530) and A. cardenasii (GKP 10017) showed field resistance to tomato
spotted wilt virus, while A. chacoensis (GKP 10602) showed no infection
either by grafting or by thrips inoculation (Subrahmanyam et al., 1985a).
Similarly Herbert and Stalker (1981) found a high level of resistance to
peanut stunt virus in species of sect. Arachis, A. duranensis (K 7988), A.
villosa (B 22585) and A. villosa var. correntina (Manfredi 8); Caulorhizae;
Erectoides; and Rhizomatosae. For peanut mottle virus, Demski and Sowell
(1981) identified six accessions of Rhizomatosae as immune, and
Subrahmanyam et al. (1985) reported no infection after mechanical or air-
brush inoculation in GK 12922, A. chacoensis (GKP 10602), A. cardenasii
(GKP 10017) and A. correntina (GKP 9530). A. monticola (GKPBSSc
30062) was found resistant to cylindrocladium black rot, (Fitzner et al.,
1985). Castillo et al. (1973) reported resistance to northern rootknot
nematodes in three accessions of Rhizomatosae, and Nelson et al. (1989,
1990) found 55 accessions of 22 species of Arachis with resistance to
rootknot nematode (Meloidogyne arenaria).
A large number of Arachis species accessions have also been screened
for resistance to insects such as thrips, leaf-hoppers and corn earworms
TABLE 4.7 Range of variation in cultivated groundnut observed at feRfSAT,
Patancheru
Character Minimum Maximum Intermediate(s)

Life form Annual
Growth habit Erect Procumbent Decumbent
Branching pattern Sequential Alternate Irregular
Stem pigmentation Absent Present
Stem hairiness Glabrous Woolly Hairy, very hairy
Reproductive branch length >lcm lOcm Continuous
Number of flowers/ 1 5 2,3,4
inflorescence
Peg colour Absent Present
Standard petal colour Yellow Garnet Lemon yellow, light
orange, orange, dark
orange
Standard petal markings Yellow Garnet Lemon yellow, light
orange, orange, dark
orange
Leaf colour Yellowish Dark green Light green, green,
green bottle green
Leaflet length 17mm 94mm Continuous
Leaflet width 7mm 52mm Continuous
Leaflet LIW ratio 1 6 Continuous
Leaflet shape Cuneate Lanceolate Obcuneate, elliptic
Hairiness of leaflet Sub glabrous Profuse and Scarce and short, scarce
long and long, profuse and
short
Number of seeds/pod S 2,3,4
Pod beak Absent Very Slight, moderate,
prominent prominent
Pod constriction Absent Very deep Slight, moderate, deep
Pod reticulation Smooth Prominent Slight, moderate
Pod length 14mm 65mm Continuous
Pod width 7mm 20mm Continuous
Seed colour pattern One Variegated
Seed colour Off white Dark purple Yellow, shades of tan,
rose, shades of red,
grey-orange, shades
of purple
Seed length 4mm 23mm Continuous
Seed width Smm 13mm Continuous
lOO-seed weight 14 g 136 g Continuous
Days to emergence 4 18 Continuous
Days to 50% flowering 17 54 Continuous
Days to maturity 75 >155 Continuous
Fresh seed dormancy o days >66 days Continuous
Oil content 31.8% 5S.0% Continuous
Protein content 15.S 34.2 Continuous
TABLE 4.8 Results of screening of Arachis germplasm at fCRfSA T Center and
Texas A & M
ICRISAT Texas A & M
Specific trait Number Number Number Number

screened identified screened identified
Disease resistance:
Early leaf spot 2500 28 (2)
Late leaf spot 9400 76 (26) 2500 40 (3)
Rust 9400 141 (35) 1500 12
TSWV 7400 23 (6)
PMV 1800 2 (2)
Aspergillus flavus 582 17 (4)
Pod rot 3222 24 (6)
Sclerotinia blight 4100 1 (1)
Wet blotch 50 35 (4)
Pest resistance:
Thrips 5000 14 (7)
lassids 6500 30 (7)
Termites 520 20 (6)
Aphids 300 4 (1)
Lcafminer 930 18 (6)
Rootknot nematodes 116 55 (3)
Multiple resistance 9400 85 (45) 2500 12 (3)
Abiotic stresses/nutrition:
Drought 742 38 (8)
N Fixation 342 4 (2)
High oil 8868 44 (10)
High protein 8868 51
Numbers in parentheses indicate number commonly used in breeding programmes.
(Stalker and Campbell, 1983), army worm (Lynch et al., 1981), lesser
cornstalk borers (Stalker et al., 1984) and spider mites (Leuck and
Hammons, 1968; Johnson et al., 1977). High levels ofresistance have been
identified in a number of accessions (Stalker and Moss, 1987).
Wild Arachis species have also been studied for nutritional quality and
some species, such as A. villosulicarpa, have been reported to have high
(1.44-1.66%) tryptophan contents (Amaya et al., 1977). Recently A. sp.
(KSSc 36008), A. paraguariensis, A. rigonii and A. appressipila have been
found with more than 60% oil content, while A. sp. GK 12922, A. sp. VKR
6110 and A. sp. GK 30126 have high linoleic acid levels (Jambunathan et
al.,1991).
Conservation of germplasm 129
4.11 CONSERVATION OF GERMPLASM
Most groundnut germplasm has been conserved in the form of either pods
or seed except for some accessions (most of which belong to sect.
Rhizomatosae) which produce very few or no seed. These non-seed pro-
ducers are being conserved as plants. The major repositories of the world
collection of groundnut germ plasm are genebanks at ICRISAT; in the
USA at the Southern Regional Plant Introduction Station in Georgia, the
North Carolina State University at Raleigh and the Texas Agricultural
Experiment Station at Stephenville; in Brazil at Empresa Brasileira De
Pesquisa Agropecuaria (EMBRAPA)/Centro Nacional de Recursos
Geneticos (CENARGEN), Brasilia and Instituto Agronomico, Campinas;
and in Argentina at Instituto Botanica del Nordeste (lEONE), Corrientes
and Instituto Nacional de Technologia Agropecuaria (INTA), Manfredi.
It is established that the two key factors influencing the seed's viability
during storage are temperature and moisture. Duration of viability is
mainly influenced by seed moisture content. Control and optimization of
these factors can dramatically improve the longevity of seeds in storage. At
ICRISAT, and probably in most other gene banks, the seeds are dried to
5-7% moisture content and are stored in controlled-environment
chambers with low temperature and humidity, following the guidelines of
the International Board for Plant Genetic Resources (IEPGR). Many
locations have three storage regimes:
1. Short-term chambers which are maintained at a temperature around

18 DC and relative humidity (RH) of 30-50%. They hold freshly har-
vested material before it is transferred to chambers with lower tempera-
ture and humidity.
2. Medium-term chambers which are maintained at 4 DC and 20-30% RH
and conserve almost all the accessions in the form of pods; made up
principally of the working and/or active collections.
3. Long-term chambers which are maintained at -20 DC with or without
control of humidity. Almost all the germplasm will eventually be stored
in these chambers as shelled seeds in closed containers (aluminium
pouches at ICRISAT) to comprise the base collection.
At ICRISAT the present emphasis is to bring all the accessions into

long-term chambers with a recommended amount of seeds. This will
minimize chances of genetic drift and loss of seed viability during conser-
vation. During transfer of germplasm to different storage conditions,
utmost care has to be taken to avoid any chances of genetic contamination
by mechanical mixing. Pathologists and virologists sample the materials to
ensure that the genebanks are stocked only with pathogen-free seeds.
Large scale multiplications, which would reduce the number of multiplica-
tion generations, can further assist in reducing chances of genetic drift or
mutations during conservation; however, low seed multiplication rate and
the large seed size of the groundnut are serious constraints.
The preservation of wild Arachis species presents special problems,
particularly in the case of accessions that produce few, if any, seeds and
these only after long periods. Rhizomatous accessions produce very few
seeds and are also quite heterozygous. Conservation of these seeds may
not represent the total variability present in the parental clones and/or
their populations. These accessions are being conserved as plants, which
are propagated vegetatively. Conservation of the wild Arachis species,
even by conventional methods, demands tremendous effort. Because of
their perennial nature, they can pose problems of disease and pest build-up
if maintained in a glasshouse. In the field, they have to be isolated to avoid
outcrossing and mechanical mixing because of their extensive vegetative
growth. Therefore most of these accessions in the USA and at ICRISAT
are maintained in containers, i.e. pots, baskets or concrete rings. Excess
vegetative growth necessitates regular maintenance of these stocks through
frequent pruning and repotting.
It is advisable to maintain these stocks of vegetatively propagated
species in duplicate and under diverse conditions so that they can adapt
and can be suitably conserved without the danger of being lost. This
practice is quite successful at present at ICRISAT, the Texas A & M
University Agricultural Experiment Station at Stephenville, the Crop
Science Department of North Carolina State University, USDA at Griffin,
Georgia, EMBRAPA/CENARGEN in Brazil and IBONE and INTA in
Argentina. This international team effort has resulted in successful germ-
plasm conservation with free exchange of material and information limited
only by the quarantine restrictions of individual countries and by financial
resources.
The research on chemical, physical and environmental factors affecting
the viability of seed during storage has been very limited as has the
utilization of high technology in vitro techniques for conservation of germ-
plasm. There are possibilities of overcoming many constraints, particularly
in the case of wild Arachis species, by using in vitro techniques for
conservation micropropagation and germplasm distribution. The vegeta-
tively propagated rhizomatous species and many other species that pro-
duce very few seeds might be stored as shoots and embryo axes in vitro or
in cryopreservation. However, before this can be attempted it will be
necessary to establish protocols for regeneration of plants from shoots and
excised embryo axes. Recently, a protocol was established at ICRISAT for
germination of embryo axes excised from healthy mature seeds (Adib et
ai., 1991) in an attempt to overcome the problem of large seed and pod
size, which restricts conservation of large numbers of seeds with the full
range of variability. The in vitro techniques could also help in overcoming
low seed multiplication through micropropagation, but suitable regener-
ation protocols from various explants causing no genetic instability and
Utilization 131
culture-induced variation are needed to make it operative on a regular
basis. Once this is achieved, the technique could be used in distribution of
germplasm, particularly of those wild Arachis species which produce very
few seeds. Strengthening of research in these areas could make utilization
of groundnut germ plasm more effective and help fill some of the existing
gaps in genebanks.
4.12 UTILIZATION
Breeding activities around the world have made significant progress in the
last two decades but exploitation of available genetic variability is still
limited. Breeders prefer their known sources of previously identified resist-
ances and are often reluctant to use those recently identified because of
lack of information on their reaction to other biotic and abiotic hazards,
centre of origin, and whether they have the same components of resistance
governed by the same genes/alleles. Poor agronomic potential of these
sources is another possible factor contributing to this situation. Hesitation
in utilizing new germplasm sources in resistance breeding is dangerous, as
the old sources may rest on a very narrow genetic base. Epiphytotics of any
pathogen may wipe out crops, causing considerable economic loss and
subsequent scarcity of planting materials. Interspecific variability has not
been fully exploited, though it has provided encouraging results (Stalker et
at., 1979; Singh, 1985b; Singh and Gibbons, 1986; Singh, 1989; ICRISAT,
]986; Simpson, 1991a).
Progress in breeding for resistance to early and late leaf spot and rust has
been significant. Southern runner, a high-yielding cultivar resistant to late
leaf spot has been released in the USA. Resistance breeding at ICRISAT
has developed high-yielding cultivars ICG(FDRS)4 and ICG(FDRS)lO
with strong resistance to rust and low level resistance/tolerance to late leaf
spot. Indian national programmes have also released several other lines.
Using several wild Arachis species from section Arachis, ICRISAT
breeders have been able to transfer disease resistance into A. hypogaea. A
large number of A. hypogaea-Iike interspecific derivatives with high yield
and resistance to rust and/or late leaf spot have been developed involving
A. cardenasii, A. batizocoi, A. duranensis, A. stenocarpa (HLK-41O, = A.
stenosperma) and A. chacoensis (ICRISAT, 1985; Singh and Gibbons,
1986). Recently, ICRISAT scientists have also identified an interspecific
line, 259-2, originating from crosses with A, cardenasii, with multiple
resistance to rust and to early and late leaf spots (Singh, 1988; ICRISA T,
1990). Resistance to groundnut rosette virus has been identified from an
interspecific derivative involving A. chacoensis (ICRISA T, 1992). Simpson
(1991a) has reported a successful pathway for introgressing leaf spot
resistance from A. cardenasii and A. chacoensis through A. batizocoi.
Introgression of nematode resistance is reported by Simpson (1991a) and
by Starr et al. (1990).
Progress in breeding for resistance to soil-borne fungi, aflatoxin develop-
ment and virus diseases such as tomato spotted wilt virus (bud necrosis) has
been very difficult and slow. Smith et al. (1991) have released a cultivar,
Tamspan 90, with high levels of resistance to pythium pod rot and scleroti-
nia blight. The research efforts in these directions have not been adequate
but there may also be a dearth of qualitative sources of resistance for most
of these diseases.
4.13 CONCLUSIONS AND PERSPECTIVES
Significant progress has been made in biosystematics of section Arachis of

the genus Arachis, leading to identification of cross-compatibility relation-
ships, genomes and phylogenetic relationships between the diploid species
of section Arachis and the cultivated tetraploid species A. hypogaea. This
information is of vital importance and has led to the establishment of
several breeding strategies for exploitation of genetic resources available in
these species. Several species have already contributed useful material.
However, progress in taxonomy and biosystematics in the genus Arachis as
a whole, particularly in relation to the seven other sections of the genus,
has been very slow. There is an urgent need to provide an appropriate
taxonomic and biosystematic treatment of the genus to avoid perpetuating
confusion and to facilitate free communication and better exchange of
information and material. This would help towards effective utilization of
the vast genetic reservoir offered by the wild Arachis species for improving
the agronomic and pathogen resistance status of A. hypogaea.
A. hypogaea is a segmental allopolyploid which originated from the
diploid species of sect. Arachis with AA and BB genomes very similar to
those present in A. hypogaea. The wild species of sect. Arachis hybridize
with A. hypogaea, and introgression of desirable genes from them is
possible through appropriate ploidy and genomic manipulations which
increase effective meiotic recombination.
With regard to groundnut genetic resources, there are positive trends in
collection characterization, conservation and utilization. Considerable pro-
gress has been made and large numbers of accessions are being conserved
in various state, national and international genebanks. However, there is
still much potential diversity and there is the urgent need to collect
additional germplasm from centres of origin and diversity in South
America. Also, areas of early introduction in Africa and Asia are in danger
of extinction and need attention. Efforts on characterization and evalu-
ation of germplasm need to be accelerated. Information on preliminary
characteristics and genetic potential is essential for utilization otherwise
these collections will continue to be underused. Special attention needs to
References 133
be given to the already identified sources of desirable traits for a thorough
evaluation to provide a complete picture of their merits and demerits, so
that they can be utilized effectively and with confidence by breeders.
Significant progress has been made in genetic improvement of the
groundnut but greater advances can be expected, particularly in the field of
breeding for disease resistance. Many sources of resistance to biotic and
abiotic stresses have been identified within the cultivated A. hypogaea but
the materials often lack desired quality characteristics. Efforts to identify
or increase variability combined with high quality characters are needed.
Considerable attention has been given to evaluation of wild Arachis
species, which have been found to possess high levels of resistance or
immunity to many diseases such as rust, early and late leaf spots, peanut
stunt virus, tomato spotted wilt virus, nematodes and other pests. Some of
the cross-compatible species within sect. Arachis have been exploited
successfully and stable 40-chromosome hybrid derivatives have been ob-
tained in several interspecific crosses. The high level of resistance of these
derivatives indicate that wild Arachis species can contribute much to
improvement of the cultivated groundnut. Species outside sect. Arachis
have not been successfully used because of differences in ploidy level,
cross-incompatibility and genomic incompatibility. Methodologies of em-
bryo rescue have not been productive and hence attention should be given
to techniques dealing with manipulation at the cellular and molecular level,
e.g. protoplast fusion and genetic transformation. However, establishment
of highly productive regeneration protocols are essential prerequisites.
These will open new vistas for exploitation of not only wild species of genus
Arachis for introgression of desirable genes but also of genes beyond
Arachis, present elsewhere in the plant kingdom.
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correntina. A. diogoi e A. villosulicarpa. Ciencia e Cultura 29.319-321.
Resslar. P.M. (1980) A review of nomenclature of the genus Arachis L. Euphytica 29.813-817.
Resslar. P.M. and Gregory. W.e. (1979). A cytological study of three diploid species of the
genus Arachis L. Journal of Heredity 70. 13-16.
Resslar. P.M .. Stucky. J.M. and Miksche. J.P. (1981) Cytophotometric determination of the
amount of DNA in Arachis L. Sect. Arachis (Leguminosae). American Journal of Botany 68.
149-153.
Seetharam. A .. Nayar. K.M.D .. Sree Kantaradhya R. et al. (1973) Cytological studies on
interspecific hybrids of Arachis hypogaea x A. duranensis. Cytologia 38.277-280.
Sharief. Y .. Rawlings. J.O. and Gregory. W.e. (1978) Estimates of leafspot resistance in three
interspecific hybrids of Arachis. Euphytica 27. 741-751.
Simpson. e.E. (1982) Collection of Arachis germplasm (1976-1982). Plant Genetic Resources
Newsletter 52. 10-12.
Simpson. e.E. (1984) Plant Exploration: Planning. Organization. and Implementation with
Special Emphasis on Arachis. in Conservation of Crop Germplasm - An International
Perspective. (ed. W.L. Brown. T.T. Chang et al.). Crop Science Society of America Special
Publication. No.8. pp. 1-20.
Simpson. e.E. (1990) Collecting wild Arachis in South America - Past and Future. in IBPGR
1990. International Crop Network Series. 2. Report of workshop in genetic resources of wild
Arachis species. Including Preliminary Descriptors for Arachis (IBPGRIlCRISA T).
International Board for Plant Genetic Resources. Rome. p. 10-37.
Simpson. e.E. (1991a) Pathways for introgression of pest resistance into Arachis hypogaea L.
Peanut Science 18. 22-26.
Simpson. e.E. (1991b) Global collaborations find and conserve the irreplaceable genetic re-
sources of wild peanut in South America. Diversity 7. 59-61.
Simpson. e.E .. Higgins. D.L.. Thomas. G.D. et al. (1992) Catalog of Pas.lport data and
minimum descriptors of Arachis hypogaea L. germplmlN collected in South America. 1977-
1986. M P-1737. Texas Agricultural Experiment Station. Texas A & M University System.
College Station. Texas. p. 244.
Singh. A.K. (1985a) Genetic introgression from compatible wild species into cultivated ground-
nut. Proceedings of International Workshop on Cytogenetics of Arachis. 31 Oct.-2 Nov. 1983.
ICRISAT. Patancheru. India. pp. 107-117.
Singh. A.K. (l985b) Cytogenetic analysis of wild species of Arachis. Project Report (1978-82).
ICRISAT. Patancheru. India. p. 71.
Singh. A.K. (1986a) Utilization of wild relatives in the genetic improvement of Arachis hypogaea
L. 7. Autotetraploid production and prospects in interspecific breeding. Theoretical and
Applied Genetics 72. 164-169.
Singh. A.K. (1986b) Utilization of wild relatives in genetic improvement of Arachis hypogaea L.
8. Synthetic amphidiploids and their importance in interspecific breeding. Theoretical and
Applied Genetics 72.433-439.
Singh. A.K. (1988) Putative genome donors of Arachis hypogaea (Fabaceae) evidence from
crosses with synthetic amphidiploids. Plant Systematics and Evolution 160. 143-151.
Singh. A.K. (1989) Exploitation of Arachis species for improvement of cultivated groundnut.
Progress Report 1988. ICRISAT. Patancheru. India. p. 73.
Singh. A.K. and Gibbons. R.W. (1986) Wild species in crop improvement: Groundnut - a case
study. in Advances in genetics and crop improvement. (eds P.K. Gupta and l.R. Bahl).
Rastogi Publication. Meerut. p. pp. 297-308.
Singh. A.K. and Moss. J.P. (1982) Utilization of wild relatives in genetic improvement of
Arachis hypogaea L. 2. Chromosome complements of species of section Arachis. Theoretical
and Applied Genetics 61.305-314.
Singh. A.K. and Moss. J.P. (1984a) Utilization of wild relatives in genetic improvement of
Arachis hypogaea L. 5. Genome analysis in section Arachis and its implication in gene
transfer. Theoretical and Applied Genetics 68. 350-364.
Singh. A.K. and Moss. J.P. (1984b) Utilization of wild relatives in genetic improvement of
Arachis hypogaea L. 6. Fertility in triploids. Cytological basis and breeding implications.
Peanut Science 11. 17-21.
Singh. A.K .• Sivaramakrishnan. S .• Mengesha. M.H. et al. (1991) Phylogenetic relations in
section Arachis based on seed protein profile. Theoretical and Applied Genetics 82.593-597.
References 137
Singh, kK., Venkateshwar, A., Moss, J.P. et al. (1990) Chromosome number and karyomor-
phology of some new accessions of genus Arachis. International Arachis Newsletter 8, 11-14.
Smartt, J. (1965) Cross-compatibility relationships between cultivated peanut Arachis hypogaea.
L. and other species of genus Arachis. PhD Thesis, North Carolina State University,
Raleigh.
Smartt, J. (1990) Grain Legumes, in Evolution and Genetic Resources, Cambridge University
Press, Cambridge, pp. 30-84.
Smartt, J. and Gregory, W.e. (1967) Interspecific cross-compatibility between the cultivated
peanut Arachis hypogaea L. and other members of the genus Arachis. O!eagineaux 22,
455-459.
Smartt, J., Gregory, W.e. and Gregory, M.P. (1978a) The genomes of Arachis hypogaea 1.
Cytogenetic studies of putative genome donors. Euphytica 27, 665-675.
Smartt, J., Gregory, W.e. and Gregory, M.P. (1987b) The genomes of Arachis hypogaea 2. The
implications in interspecific breeding. Euphytica 25, 677-680.
Smith, 0.0., Simpson, e.E., Grichar, W.J. and Melouk, H.A. (1991) Registration of 'Tamspan
90' peanut. Crop Science 31, 1711.
Stalker, H.T. (1981) Intersectional hybrids in the genus Arachis between section Erectoides and
Arachis. Crop Science 21,359-362.
Stalker, H.T. (1991) A new species in section Arachis of peanut with a 0 genome. American
Journal of Botany 78,630--637.
Stalker, H.T. and Moss, J.P. (1987) Speciation, cytogenetics and utilization of Arachis species.
Advances in Agronomy 41, 1-40.
Stalker, H.T. and Dalmacio, R.D. (1981) Chromosomes of Arachis species, Section Arachis
(Leguminosae). Journal of Heredity 72,403-408.
Stalker, H.T. and Campbell, W.V. (1983) Resistance of wild species of peanut to an insect
complex. Peanut Science 10, 30-33.
Stalker, H.T. and Wynne, J.e. (1979) Cytology of interspecific hybrids in the section Arachis of
peanuts. Peanut Science 6, 110-114.
Stalker, H.T., Wynne, J.e. and Company, M. (1979) Variation in progenies of an Arachis
hypogaea x diploid wild species hybrid. Euphytica 28, 675--684.
Stalker, H.T., CampbelL W.V. and Wynne, J.e. (1984) Evaluation of cultivated and wild
peanut species for resistance to the lesser cornstalk borer (Lepidoptera: Pyralidae) Journal of
Ecological Entomology 77,53-57.
Starr, J.L., Schuster, G.L. and Simpson, e.E. (1990) Characterization of the resistance to
Meloidogyne arenaria in an interspecific Arachis spp. hybrid. Peanut Science 17, 106-108.
Subrahmanyam, P., Ghanekar, A.M., Nolt, B. et al. (1985a) Resistance to groundnut diseases in
wild Arachis species, in Proceedings of International Workshop on Cytogenetics of Arachis,
31 Oct. 2 :--Iov. 1983, ICRISAT, Patancheru, India, pp. 49-55.
Subrahmanyam, P., Moss, J.P. and Rao, V.R. (1983) Resistance to peanut rust in wild Arachis
species. Plant Disease 67, 209-212.
Subrahmanyam, P., Moss, J.P., and McDonald, D. et al. (1985b) Resistance to leafspot caused
by Cercosporidium personatum in wild Arachis species. Plant Disease 69,951-954.
Subrahmanyam, P., McDonald, D., Redd, L.J. et al. (1990) Resistance to rust and late leafspot
of groundnut at ICRISAT Center. Problem and Progress. Proceedings of Fourth Regional
Groundnut Workshop for South Africa, 19-23 Mar. 1990, Arusha, Tanzania. ICRISAT,
Patancheru, India, pp. 85-92.
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Prant!) TeiL Abt. 3: 70--388 (Arachis). Verlag von Wilhelm Engelman, Leipzig.
Tombs, M.D. and Lowe, M. (1967) A determination of the sub-units of arachin by osmometry.
Biochemical JournalI8t, 181-187.
Valls, J.F.M., Rao, V.R., Simpson, e.E. et al. (1985) Current status of collection and conser-
vation of South American groundnut germplasm with emphasis on wild species of Arachis.
Proceedings of an International Workshop on Cytogenetics of Arachis, 31 Oct.-2 Nov. 1983,
ICRISAT, Patancheru, India, pp. 15-39.
Williams, E.D. (1991) Exploration of Amazonian Bolivia yields rare peanut landraces. Diversity
5, 12-13.
Wynne, J.e. and Halward, T. (1989) Cytogenetics and genetics of Arachis, in Critical Reviews in
Plant Science, (ed. B.V. Conger) CRC Press, Boca Raton, pp. 189-220.
CHAPTER 5
Reproductive biology and

development
Peter Coolbear
The possibility that any plant - never mind a major commercial crop -
might possess a reproductive strategy whereby fertilized ovaries bury
themselves in the soil before seed development can properly commence is,
to say the least, unusual. It is understandable, therefore, that the repro-
ductive botany of the groundnut (peanut) remained a source of surprise
and confusion well into the middle of this century and that it was not until
then that a truly definitive description of this process was published (Smith,
1950). Forty years later, this geocarpic habit still raises a series of intriguing
and fundamental questions for seed scientists.
The aim of this chapter is to review the major events in this process from
flower development onwards. Special emphasis has been placed on factors
affecting seed yield and quality, taking the quality aspects of the discussion
through to seed storage and the particular problems of groundnut which
result from the fact that 50% of the weight of the seed may be oil reserve.
5.1 FLOWER DEVELOPMENT, POLLINATION AND

FERTILIZATION
Groundnuts may flower as early as 3 weeks after planting. Van Rossem

and Bolhuis (1954) found floral initiation in the cotyledon axillary buds at a
very early stage in the seedling growth of a spanish cultivar, Schwarz 2l.
Typical flowering patterns for both a virginia and a spanish cultivar are
shown in Figure 5.1. In this study, flowering occurred over a period of
about 11 weeks, with peak production occurring at around 8 weeks after
sowing in the spanish type and after 10 or 11 weeks in the more prolifically
flowering virginia type. In both cultivars flowering was clearly cyclical and
this pattern was apparently independent of both day length and environ-
Flower development, pollination and fertilization 139
~ 25 ,---------------------------------------------------,
-c Virginia Jumbo Runner
....
8. e-------e Spanish type
20
"8
:J
-c
ea. 15
-oJ
r::::
c
a.
~ 10
a.
5 6 7 8 9 10 11 12 13 14 15 16
Weeks from sowing
Figure 5.1 Flowering pattern in groundnut. Typical patterns of flower production

in two cultivars grown in North Carolina, USA. (Redrawn from Smith, 1954.)
ment per se. Bagnall and King (1991 a,b) have recently demonstrated for a
range of cultivars that while the onset of flowering is independent of
daylength (except under low light and temperature conditions when short
days are promotory), the number of flowers produced is greatly enhanced
by short days. Flowering rates are also a function of plant size and are
promoted by increasing mean temperatures up to about 30°C and/or
increased light intensity. On the other hand, they may be reduced by
intraplant competition with developing fruits and environmental stresses of
various kinds, in particular a shortage of nutrients (Smith, 1954).
The compressed spical inflorescences of Arachis hypogaea comprise at
least three flowers but it is unusual for more than one to elongate at the
same time, although the second and third flower may then expand on
successive days in warm weather. In cool or wet conditions this interval
may be greatly increased. Opening usually takes place in the early morn-
ing; on the day prior to this, the sessile 6-10 mm flower bud will double in
length, after which the rate of expansion growth of the hypanthium more
than trebles overnight. At anthesis the flower may be up to 70 mm in length
from the base of the hypanthium, a 20 mm standard being borne on a
50 mm hypanthial tube (Figure 3.5). During this time the style elongates in
pace with the hypanthium and remains close to the dehiscing anthers.
Self-pollination normally occurs either just before or at the same time as
flower opening, although it should be noted that some wild species of
groundnut, (e.g. A. lignosa) may require insect pollination (Banks, 1990).
140 Reproductive biology and development
Generally, in A. hypogaea, groundnut ovaries contain between two and
five ovules, most commonly two (Figure 5.2(a». However, development of
only a single seed in the fruit is frequent, as is the production of 'pops' -
empty pods. In certain conditions, losses of potential yield can be more
than 90% (Smith, 1946). The reasons for this will be explored later.
Sporogenesis and gametogenesis occur 2 days prior to anthesis when bud
length is around 5 mm (Smith, 1956a). There is general agreement in the
literature that the antipodal cells degenerate several hours before fertiliz-
ation takes place (Smith, 1956a, Periasamy and Sampoornam, 1984; Pattee
and Mohapatra, 1987). This is in marked contrast to many other legumes
where these cells persist for some time (usually days) after fertilization and
are thought to playa key role in initial endosperm development, possibly
by providing a readily available supply of specific nutrients or endogenous
plant growth regulators as they break down (e.g. Chen and Gibson, 1971).
A well established feature of the embryo sac is that it contains a large
quantity of starch grains which will provide the nutrients for early endo-
sperm growth (Reed, 1924; Smith, 1956b). Details of embryo sac anatomy
are shown in Figure 5.2(b).
Considering the distance it has to grow, pollen tube extension is rapid.
Germination of pollen is promoted by red light, auxin, gibberellic acid
(GA) and many of the phenols present as glycoside derivatives in the
stigma and style. Postgerminative pollen tube growth requires both tran-
scriptional and translational activity and is promoted by ethylene and red
light, although GA can substitute for the phytochrome response in dark-
ness (Malik and Chhabra, 1978). Smith (1956a) estimated that, on average,
a pollen tube took about 9 h to reach the ovary and fertilization occurred
10-18 h after pollination. Later authors dispute his view that the synergids
disintegrate prior to this event. Periasamy and Sampoornam (1984) suggest
that the cellular synergids are essential to this process and that pollen tube
penetration actually occurs through the vacuole of one of them, although
Pattee and Mohapatra (1987) were unable to confirm this. In Smith's
(1956a) study less than 7% of ovules failed to be fertilized, although there
was a small additional percentage where there was successful fusion of only
one of the two pollen nuclei. Malik and Chhabra (1978), however, cite very
much lower fertilization percentages in India. In their survey, successful
pollination is quoted as being between 5% and 59% in spreading types and
22-68% in bunch forms with extracted pollen having very poor viability. It
is unclear whether this difference between India and the USA is a function
of local varieties or environment.
S.2 FRUIT DEVELOPMENT
After fertilization the flower withers rapidly and over a period of several
days the fruit develops as a positively geotropic stalk-like structure, the
Fruit development 141
I I (b)
(a)
I I
I I
I I
/ I
I I
I J
\
"; f---f 0.01 mm
.....-.... 0.1 rnm

es
~O.lmm t----I 0.1 mm
1-------11 0.1 mm
Figure 5.2 Embryology of groundnut (a, apical ovule; b, basal ovule; c, cytoplas-
mic strands; da, degenerating antipodals; ds, degenerating synergids; dsu, degener-
ating suspensor; e, embryo sac; ea, egg apparatus; f, funiculus of apical ovule; h,
hypanthium; i, integumentary tapetum; m, meristem; pe, pro-embryo; s, synergids;
sg, starch grains; su, suspensor): (a) an ovary at anthesis; (b) embryo sac at
anthesis, with plentiful starch grains enclosed in compartments defined by cytoplas-
mic strands; (c) four-cell stage in embryogenesis; (d) heart-shaped embryo stage;
(e) embryo after cotyledon initiation. The endosperm tissue remains poorly differ-
entiated, parts of it remaining acellular throughout development. ((a)-(d) are
interpretations of plates in Pattee and Mohapatra, 1987; (e) in Peri as amy and
Sampoornam, 1984.)
'peg', which eventually penetrates the soil. The peg is often described,
even in recent literature (e.g. Ziv 1981; Amoroso and Amoroso, 1988), as
a gynophore. As Smith (1950) points out, this is technically incorrect
because the ovary is sessile: there is no ovary stalk to elongate. The cells
which elongate comprise the basal tissue of the ovary itself and thus a more
appropriate term for the peg is a carpophore (Smartt, 1976).
Growth of the carpophore is intimately associated with embryo develop-
ment. Peg extension is slow at first, taking 5-6 days to penetrate the bracts
(Smith, 1950). As the fruit reaches 3-4 mm in length it becomes positively
geotropic and starts to grow towards the soil (Peri as amy and Sampoornam,
1984), the rate of elongation increasing rapidly between 5 and 10 days after
fertilization (Figure 5.3). By the time this rapid elongation starts, the
embryo, which is now in the 8-12 cell stage, becomes quiescent (Smith,
1956b). Reduction in embryo growth may simply be a function of nutrient
deficiency due to the high demand of the rapidly growing carpophore, or
there may be more subtle control via pH changes within the ovary apex
(Schenk, 1961). Zamski and Ziv (1976) have demonstrated that initial
proembryo development is an essential prerequisite for peg elongation and
suggest that auxin and GA produced by the proembryo promote its
growth. Ziv (1981) provides evidence which indicates that carpophore
growth is also mediated by phytochrome. The carpophore is actively
photosynthetic, containing two or three layers of chlorenchyma with abun-
dant stomata on the endodermis, these later being replaced by lenticels as
secondary thickening occurs.
Growth usually continues until penetration of the soil occurs 8-14 days
after fertilization (Schenk, 1961). Zamski and Ziv (1976) found that, in the
virginia runner type of groundnut, light inhibited pod formation and pegs
failing to reach soil might grow up to 20 cm before withering and dying.
They were also able to demonstrate that the mechanical stimulus of the peg
entering the soil is important for successful pod development, stimulating
ovary wall thickening and the diageotropic orientation of the pod due to
rapid cell enlargement on the dorsal side of the carpel near the basal ovule.
Once pod development starts, seed development resumes. If an ovule is
unfertilized there is generally little pod development around that ovule
(Periasamy and Sampoornam, 1984). Typically, basal ovules resume
growth 1-2 days before apical ones and endosperm growth recommences
1-2 days before embryo growth (Smith, 1956b; Pattee and Mohapatra,
1987) (Figure 5.3).
Depth of pod placement depends on soil structure (due to both its
mechanical properties and light penetration), but usually pods are placed
about 5 cm below the soil surface. Periasamy and Sampoornam (1984)
noted that the presence of light may not be completely inhibitory to pod
development in all cultivars. In the erect valencia groundnut some aerial
pod development does occur and viable, small seeds may be produced.
This is unusual and in general the successful development of high quality
seed is dependent upon soil penetration by the peg.
Schenk (1961) undertook a detailed study of the underground develop-
ment of whole fruits in both a virginia bunch type and a spanish type. Fresh
weights of whole pods increased very rapidly during the first 2 weeks of
subterranean growth and they attained maximum size after 3 weeks.
Subsequently there was little increase in total fresh weight, although it
fluctuated with soil moisture status. Dry weights increased more gradually,
reaching a maximum after 8 weeks in virginia bunch (Figure 5.4) and after
Fruit development 143
6 weeks in the faster developing spanish type. Respiration rates were
highest during the period of maximum pod growth (Figure 5.4). After 2-3
weeks there was very little change in shell dry weights, although fresh
weights decreased rapidly as tissues collapsed and structural changes
occurred as the shells hardened. Hull colour changes were a useful indi-
cator of pod maturity (section 5.4.2). The seeds themselves tended to
develop more slowly, following a typical pattern which will be discussed in
detail in the next section. In this study, spanish pods were ready for lifting
after about 7 weeks of underground development, and virginia bunch pods
after 11 weeks. Times vary depending on cultivar and production con-
ditions but, in general, runner types take about 180 days from planting to
80 Peg length
~.><
X"~
Pods begin to /
./
,60 penetrate soil ./
)<
E
.s
/
/
/ Onset of
.r:::
0,40 / pod development
c: ~
~ /
Ol
OJ X
.I
c.. 20 /
.X·
>l
/
..>(/><
0 .""*".~'
(72.3)
Embryo cell numbers
!!J.
Qi
() 20 Basal
'0 endosperm
Q;
.0 • •
E 10
::::J
z 0 Apical
endosperm
0
(198.8)
Endosperm growth
r
I
50
'iii
(3
::::J
c: Basal
'0 30 endosperm
Q; •
.0
E 0 •
0 0 0
::::J
z 10 Apical
endosperm
0
0 2 4 6 8 10 12 14 16
Days after pollination
Figure 5.3 The growth of peg, embryo and endosperm of cv. Virginia Runner
during peg elongation and soil penetration. (Data from Smith, 1956b.)
maturity in the US, compared with 130-135 days for virginia bunch and
around 120 days for spanish types (Woodroof, 1973).
At digging, pods may still have a moisture content of 35-55% and are
usually left to dry in the field until kernel moistures are down to 18-25%,
when they are brought in and artificially dried for storage. Late digging and
prolonged windrowing before combining both result in potential loss of
yield as pods become detached from the carpophore (Schenk, 1961;
Woodroof, 1973).
5.3 SEED DEVELOPMENT
In most orthodox seeds, whether endospermic or non-endospermic, there

are three clearly defined phases of development (e.g. Hyde et al., 1959). A
phase of rapid growth with concomitant increases in both fresh and dry
weight (Stage I) is followed by a period of food reserve accumulation,
where the rate of dry weight increase is often constant but seed moisture
contents begin to fall (Stage II). The seed then loses water in the third,
ripening stage, usually with little change in dry weight (Stage III).
These three stages summarize the different morphological and physio-
logical processes which occur during development and provide useful
reference points for subsequent discussion. During Stage I, growth is
• Whole fruit
•
dry weight
600 0 1.5
I
. 500
Maximum
dry weight
kernel ~
.
..r::. Whole fruit attained :!:
O2 uptake of
:!:
400 1.0 ~
of ..r::.
~
~ ..-
.,
~
300
..r::.
0>
.ij;
.Ea. ~
:::l ~
0
0" 200 1.5
100
I
Maximum shell dry
weight attained
0
0 2 4 6 8 10 12
TIme underground (weeks)
Figure 5.4 Oxygen uptake and dry weight increases by whole fruits of virginia
bunch groundnuts during development. (Redrawn from Schenk, 1961.)
Seed development 145
mainly by cell division. At the end of this stage all the key structures of the
seed are present, even if not fully differentiated. Growth in Stage II is thus
by cell enlargement as seed reserves are laid down. The loss of moisture
content reflects the increasing proportion of insoluble food reserves in
storage tissue. At the end of this period, maximum seed dry weight is
attained. This point has been variously referred to as morphological
(Anderson, 1944), functional (Grabe, 1956) or physiological maturity
(Shaw and Loomis, 1950). It is perhaps unfortunate that the last of these
terms is the one in general use, as the physiological development of the
seed may be far from complete at this time. Many seeds require the
stimulus of further drying to become fully germinable (e.g. Kermode et al.,
1986). It is generally accepted, however, that a seed's maximum vigour
potential (i.e. its potential to produce a strong and healthy seedling under
variously stressful field conditions) is in place at this time.
Because of the obvious difficulties of working with an indeterminate
crop with the added complication of fruits developing underground, few
workers have characterized the growth of groundnut kernels in these
terms. A study of this type, using tagged carpophores just before they
entered the soil, was undertaken by Pickett (1950) but he estimated that
there were variations in physiological age within samples of up to 7 days.
Data shown in Figure 5.5 were obtained by Aldana et al. (1972) by
harvesting only fruits produced by the second internode of the first lateral
reproductive branch of field grown plants of cv. NC-2 (a virginia bunch
variety). Although data collection did not commence until 4 weeks after
the buried ovary had become diageotropic (at which stage the endosperm
has been fully absorbed), it is possible to identify the general pattern of
seed development outlined above. For these seeds, stage I was completed
8-9 weeks after flowering when the increase in dry weight became linear.
Maximum dry weight was achieved at 13-14 weeks. At 14 weeks after
flowering, these fruits were considered ready for Hfting, at a time when
Stage III of seed development had barely begun. On inverting, seed
moisture contents are usually higher than 35% (Woodroof, 1973) and
Stage III is completed during windrowing. Depending on cultivar and local
environment, Stages I and II of seed development may be completed 8-14
weeks after flowering.
Although the above description summarizes the broad pattern of seed
development, it should be remembered that each component - seed coat,
endosperm and embryo - are genetically different and each follows its own
programme. The details of the developmental pattern for each system are
discussed individually below.
5.3.1 Embryology
After fertilization the contents of the zygote become much denser and the
cell enlarges. In common with most species, groundnut cell division lags
behind that of the endosperm, the first mitosis not occurring until the
beginning of the second day after fertilization (Smith, 1956b). The basal
cell undergoes a second mitosis within 24h but another day may elapse
before the terminal cell divides. However, as mentioned previously, after
5 days the quiescent period for both embryo and endosperm growth occurs
with rapid extension growth of the peg. Smith (1956b) found growth
resuming 10-12 days after pollination (Figure 5.3). Having observed the
contraction of the embryo sac during the quiescent phase, Pattee and
Mohapatra (1987) concluded that the system was under considerable meta-
bolic stress at this time. This suggestion tallies with the extensive losses in
potential yield which occur during pegging (section 5.6.1).
The suspensor of the developing groundnut seed is typically large. Early
growth of this organ is characterized as much by cell enlargement as by cell
division, while the cells derived from the terminal cell will continue to
divide without much expansion to produce a tight ball. After 3-4 weeks,
the undifferentiated globular embryo may consist of 500-1000 cells. At this
time the suspensor is likely to have around one tenth of this number of
1600 FWT0 ao
SMC:
I
x,.,.·)(\ I 0
I
\. I
.........
C7I 1200 X.I
60
E
........ ix
I
..
I ~
I U
.,
+'
::::E
..r:::.
. C7I aoo I
I Ul
~
40 ~
-0
Q)
Ul
Q)
400 20
0 0
6
I
a:
10 12 :14 I
16
Weeks after flowering
I I
Stage I I Stage II I Stage III

-:-I
-I-
Figure 5.5 Changes in seed weight (less testa) and seed moisture content (SMC)
during groundnut seed development. Material for this study was harvested from
fruits produced at the second internode of the first reproductive lateral in var.
NC-2. The characteristic stages of seed development are indicated on the graph.
(Calculated from Aldana et ai., 1972.)
Seed development 147
800 Cotyledons
600
400
." 200
I:
"
i;
Q.
'"
E Axis
20
10
0
6 8 10 12 14 16
Figure 5.6 Dry weight changes of the embryo axis and cotyledons during seed
development in groundnut var. NC-2. (Redrawn from Aldana et ai., 1972.)
cells; in contrast, there may now be 5000 nuclei in the acellular endosperm
(Smith, 1956b). From this stage onwards the embryo will successively
become heart -shaped (Figure 5 .2( d)) and differentiate well defined cotyle-
donary outgrowths which arise at opposite sides of the proembryo (Figure
5.2(e)). Being constrained by the embryo sac and the developing seed coat,
the cotyledons curve round in such a way that they come together in
parallel. Between them lies the plumule, which is a complex meristematic
region showing not only a terminal bud but also two lateral buds in the axes
of the cotyledons and several well developed leaf primordia (Reed, 1924;
Gelmond, 1971a; Periasamy and Sampoornam, 1984). The radicle and
hypocotyl are discernible from the suspensor region only at a late stage
(Reed, 1924) and protrude from the base of the cotyledons, making the
seed highly susceptible to mechanical damage (Gelmond, 1971a). Figure
5.6 shows how weight changes in axis and cotyledons differ as development
proceeds. Most of the dry weight of the axis has been attained by the end of
Stage I, while the bulk of cotyledon growth is in Stage II, reflecting the
storage function of this tissue. Schenk (1961) noted that seeds of virginia
bunch groundnuts were capable of germination about 6 weeks after the end
of Stage I.
5.3.2 Endosperm development

The triploid endosperm nucleus begins to divide within a few hours of
fertilization (Smith, 1956b) and the two daughter nuclei migrate to
opposite poles of the embryo sac. Each nucleus takes half the starch grains
with them (Periasamy and Sampoornam, 1984). Despite early descriptions,
subsequent endosperm development is entirely coenocytic. After 4 days
the endosperm reaches the 16 nuclei stage, by which time the starch grains
of the embryo sac had disappeared in the virginia and valencia cultivars
described by Smith (1956b) and Periasamy and Sampoornam (1984), but
they have recently been observed to persist much longer in the Indian
cultivar NS-5 (Suryakumari et ai, 1989).
As the embryo sac elongates, the endosperm remains as strands
of largely peripheral multinucleate cytoplasm, although another film of
endospermic cytoplasm surrounds the embryo (Figure 5.2(c)-(e)).
Cellularization of the endosperm does not begin until 5 or more weeks
from fertilization, when the emhryo has begun to develop its cotyledonary
outgrowths. However, there still seems to be dehate in the literature
concerning the extent of the cellularization process. Periasamy and
Sampoornam (1984) consider that cellularization only occurs at the chala-
zal end of the embryo sac, but both Prakash (1960) and Suryakumari et ai.
(1989) report endosperm cellularization right to the suspensor. Periasamy
and Sampoornam (1984) suggest the possibility of confusion of endosperm
tissue with the inner layers of the developing seed coat and the light and
scanning electron micrographs (SEMs) of Pattee and Mohapatra (1987)
would appear to bear out this suggestion. Nevertheless some SEM pictures
from these authors seem to indicate some cell wall formation near the
embryo, although they themselves refrain from speculation, recognizing
the problems of fixation and shrinkage in this fragile tissue. It is quite
likely, too, that there is some genotypic variation between cultivars. For
instance, Suryakumari et ai (1989) described the appearance of endosperm
nodules at an early stage of embryo development. These were discrete
structures within the endosperm containing four to five nuclei. There
appears to be no mention of this type of structure in other descriptions.
Whatever the case, it seems clear that cellularization of the endosperm is
incomplete and, in common with many other legumes, this tissue is wholly
absorbed as the cotyledons enlarge in Stage II of development.
5.3.3 Seed coat

Although the coat of the mature dry seed is thin and papery, it may
comprise as many as 25 layers of collapsed parenchymatous cells. There is
a considerable divergence of opinion about the origin of these layers.
Suryakumari et ai. (1989) suggest that they are entirely derived from the
outer integument; however, Periasamy and Sampoornam (1984) suggest
that the major part of the seed coat is chalazal in origin. Unfortunately the
photographic evidence for this is inconclusive in their paper while none is
presented by Suryakumari et al. (1989). While this type of development has
The biochemistry of seed development 149
been recognized in other families, it does not seem to have been reported
previously in the Leguminosae (Corner, 1976).
Schenk (1961) demonstrated that seed coats attain maximum dry weight
by the end of Stage I of development. However, their moisture contents
and metabolic activity are still high. During Stage II skins begin to darken
and tannin contents increase considerably as dry weights start to decrease.
About 8-10 weeks after fertilization, they lose moisture and collapse.
Interestingly the progress of seed coat development in the virginia bunch
type followed a very similar time course to the spanish type, despite the
fact that kernel development was so much more rapid in the latter cultivar.
There is an extensive network of eight to ten bundles of vascular tissue
through the seed coat (Periasamy and Sampoornam, 1984). This network is
crucial in supplying nutrients to the developing embryo during Stage II of
development. Schenk (1961) notes that the vascular strands in the skins of
virginia bunch types persist after the rest of the testa tissue has collapsed
around the middle of Stage II. There are, of course, no direct vascular
connections to the embryo, nutrients being unloaded into the apoplast
surrounding the developing cotyledons.
5.4 THE BIOCHEMISTRY OF SEED DEVELOPMENT
Figure 5.7 shows the patterns of DNA, RNA and protein accumulation in
cotyledons and axes of developing kernels described by Aldana et ai.
(1972). DNA levels reached a maximum in both tissues after 8 weeks of
development in these seeds, confirming that embryo growth by cell division
was essentially complete at the end of Stage I of seed development. The
loss of RNA from the cotyledons between weeks 10 and 13 (Figure 5.7 (b ))
remains unexplained. Levels of ribonuclease, although variable, were high
at all stages of development. The observation of an increase in RNA levels
after the seed was ready for harvesting was complemented in later work by
Pattee et ai. (1981), who found a dramatic fall in cotyledonary ribose as
seed matured. It is possible that these late changes may represent the onset
of germinative activity in an over-mature seed, although actual radicle
emergence is likely to be inhibited by dormancy in most cases (section 5.5).
As shown in Figure 5.6, the cotyledonary reserves accumulate during
Stage II of development. The mature seed comprises approximately 30%
protein, around 50% lipid and 12% starch (Bewley and Black, 1978).
Pickett (1950) determined the time course of accumulation of these com-
ponents in virginia bunch type (Figure 5.8), although his data may be
confounded by high variation in maturity classes at each harvest.
Seed storage protein is rich in arginine, aspartate, glutamate and glycine,
but methionine levels are low (Yatsu and Jacks, 1972). There are particu-
larly high levels of free glutamate during development, while the high
arginine levels found in immature kernels rapidly decrease as the seed
Cotyledons
200
100
·x0
OJ
...0 0
Cotyledons
.5 800 (b) RNA
a.
c:
0
]600
~
"
8.400
s: Axis
200
0
(e) Protein
...
.[ 30
600
c: .,
]
0 Axis ·x0
~20 400 ...a.
Q)
"... 01
Q)
a. :t
01 10 200
E
0 0
6 8 10 12 14 16
Figure 5.7 Accumulation of DNA, RNA and protein in axes and cotyledons
during development of groundnu,t kernels, vaT. NC-2. Note change of scale for
different tissues in graph (c). (Redrawn from Aldana et al., 1972.)
approaches maturity (Mason et al., 1969; Young, 1973). Major storage

proteins comprise a 12-14 S legumin type glycoprotein (molecular weight
=350 kD), two or more vicilins (mwt between 140 and 190 kD) and small
amounts of a 2S globulin (mwt =20 kD) (Derbyshire et al., 1974).
The pattern of protein accumulation found by Aldana et al. (1972) in a
similar type (Figure 5.7(c)) represents a study from more homogeneous
material than that used by Pickett (1950) and shows some interesting
differences from the pattern shown in Figure 5.8. Data from the study by
Aldana et al., (1972) show a triphasic pattern of protein accumulation in
the cotyledons. This is similar to that found in maize endosperm by Ingle et
The biochemistry of seed development 151
Oil •
250
200
] 150 Protein
8.
E 100
50
40
8. 20
E
Sucrose
10
o
4 10 12
Figure 5.8 Changes in the chemical composition of whole kernels during the
development of virginia bunch type groundnuts. (Redrawn from Pickett, 1950.)
al. (1965) where a lag phase in net protein accumulation during early Stage
II resulted from the interconversion (via breakdown and reassembly) of
proteins no longer required for cell division to storage proteins. If this
figure is compared with the changes in dry weight occurring in the same
seeds (Figure 5.5), it can be seen that extensive protein accumulation is still
occurring in early Stage III of seed development, i.e. after the seed has
reached maximum dry weight.
The unusually high lipid content of mature groundnut kernels is made up
largely of triglycerides whose major fatty acid components are 18:1, oleic
(=50%) and 18:2, linoleic (=24%) acids, but 16:0 through to 24:0 satu-
rated fatty acids are also present in significant amounts, as is 20: 1 un-
saturated fatty acid (Worthington et al., 1972; Bewley and Black, 1978).
Variation between genotypes is considerable: for example, in their survey
of 82 lines, Worthington et al. (1972) found oleic acid contents varied
between 36% and 69% and linoleic acid between 14% and 40%. Linolenic
acid (18:3) is usually present in only small amounts (between 0.04% and
0.5% - Pattee et al., 1970 and Wallerstein et al., 1989, respectively).
Environmental factors during seed production may also affect the oil
content of seeds (section 5.7.1).
5.4.1 The metabolism of seed development
Little work seems to have been done on the cellular biochemistry of the
seed during development. Both protein and lipid are stored in membrane-
bound bodies within the cotyledons. It is likely that the site of both lipid
accumulation and storage protein synthesis is the endoplasmic reticulum,
with dictyosomes being responsible for the accumulation of storage protein
in vacuoles. However, the details of these processes have yet to be worked
out (Bewley and Black, 1978). Yatsu and Jacks (1972) made a good case
suggesting that the lipid-containing spherosomes in groundnut and other
seeds are bounded by half unit membranes. More recently, Cao and Huang
(1986) have demonstrated that diacylglycerol acyltransferase (an enzyme
which would be required for triacylglyceride assembly in the lipid bodi.es) is
localized in the rough endoplasmic reticulum of maize and probably other
seeds. This suggests that, during lipid body development, triacylglycerides
are inserted into the middle of the lipid bilayer of endoplasmic reticulum
membranes.
Most studies on groundnut seed development have, understandably,
focused on the flavour producing compounds in the kernel. Unfortunately,
these data have often been obtained from mixed samples of pods of varying
maturity and cannot be related precisely to stages of development post-
anthesis. Thus Mason et al. (1969) demonstrated that, while many free
amino acids remained relatively constant during development, there was a
slow but steady increase in phenylalanine as seeds matured and, as already
mentioned, there were dramatic decreases in arginine levels such that the
amount in the kernel had fallen approximately ten-fold from its levels in
early development. Similarly, they demonstrated that sucrose levels fell
during development, then rose again as seeds finally matured, confirming
earlier findings by Pickett, 1950 (Figure 5.7(b)). Sucrose is by far the most
predominant soluble carbohydrate, comprising about 80% of the total at
maturity, with stachyose forming another 16% (Pattee et al., 1981). Starch
levels peak around the end of Stage I of seed development and then begin
to fall (Figure 5.7(b)). A high proportion of the starch during early
development is present in the seed coat (Schenk, 1961; Suryakumari et al.
1989).
Research on volatiles present in developing groundnut seeds by Pattee
et al. (1970) indicate that the maximum metabolic activity of seeds occurs
during Stage II of seed development and is probably related to larger seed
size while moisture contents are still adequate, rather than high levels of
metabolic activity per se. Schenk's (1961) data for pod respiration show
that this reaches a maximum around the beginning of Stage II of seed
development (Figure 5.4). During active lipid synthesis the respiratory
quotient of developing kernels approaches two, falling to unity as this
process stops. One of the most noteworthy aspects of the study by Pattee et
al. (1970) was the high level of lipoxygenase activity detected from the
Seed dormancy 153
ninth week of development onwards: the activity of this enzyme might be
expected to reduce the levels of polyunsaturated fatty acids accumulated
by the maturing seed. Levels of conserved lipoxygenase in dried kernels
may be important in the subsequent storage behaviour of seeds (section
5.8.2).
5.4.2 Maturity indices based on the process of development

A major problem in groundnut production is determining the appropriate
time to harvest. Inevitably, because of the plants' extended flowering
pattern, there will be a wide range of maturity classes within the crop.
Immature kernels tend to produce off-flavours, especially when cured at
high temperatures (e.g. Kramer et al., 1963). Various attempts have been
made to develop rapid methods of determining kernel maturity, some of
which are based on biochemical changes occurring during development.
These include spectrophotometric analysis of whole seeds to screen for the
high xanthophyll levels found in immature cotyledons (Kramer et al.,
1963); assays of volatiles produced by seeds, the levels of which decrease as
seeds mature (Pattee et al., 1970); density measurements of cured kernels
which decrease as they mature, due to the fact that tissues of mature seeds
are less likely to collapse during heating (Miller and Burns, 1971);
or screening for the high arginine levels found in immature seeds
(Young, 1973). However, the most useful method seems to be based on
measuring the darkening of internal hull tissues during maturation
(e.g. Miller and Burns, 1971). It has recently been shown that this
browning is due to flavonoid metabolism in the maturing pod, especially
an increase in the pigment luteolin. As such changes are not normally
affected by variations in environment during seed production, this simple
method of maturity determination is more reliable than most (Daigle et al.,
1988).
5.5 SEED DORMANCY
Dormancy may occur in many varieties of groundnuts and is found in both

bunch and runner types (Toole et al., 1964). In general the presence of
dormancy in groundnut is not a practical problem as it will usually wear off
in storage before the next planting season. However, lack of dormancy in
some lines can cause vivipary in areas when rains delay harvest, e.g.
Maharashtra in India (Bhapkar et al., 1986) and parts of Africa (Smartt,
1976).
Various authors (e.g. review by Bhapkar et al., 1986) have demonstrated
that the coat-imposed component of groundnut seed dormancy involves
water soluble germination inhibitors and, in some cases, resistance to
initial water uptake by the dry seed. Sengupta (1989) has suggested that the
inability to leach phenolic compounds rapidly on imbibition may be one
cause of seed dormancy. Despite the assertions of Bhapkar et ai. (1986),
ground nut dormancy mechanisms do not reside entirely in the seed coat:
there is also a component of 'true', embryo-based dormancy. Toole et ai.
(1964) clearly demonstrated that, whilst seed coat removal improves
germinability of many maturity classes, ethylene may be required to ob-
tain full germination of either mature intact seeds or immature decoated
ones. In a single lot varietal comparison, they demonstrated that there
may be considerable differences in dormancy between varieties, but it is
equally clear that different lots of the same variety may show differing
dormancy characteristics depending on production conditions. For
example, dormant seeds may lose dormancy if digging is delayed and
they are allowed to remain in the soil after maturity. As a general rule,
immature seeds of a dormant line have deeper dormancy than mature
ones, while basal seeds are more dormant than apical seeds in the same
maturity class.
In a series of papers, Ketring and Morgan (1970, 1971, 1972) have
investigated the embryo component of groundnut seed dormancy in detail.
They have shown that the evolution of ethylene by the embryo is an
essential prerequisite for germination and have estimated that groundnut
seeds must be able to produce 2-3 nl/g fresh weight in order to break
dormancy, the required threshold ethylene concentration for basal seeds
often being twice that of apical seeds. The germination promoting effects
of GA or, more potently, cytokinin are via their promotion of ethylene
production. Surprisingly, auxin, known to promote ethylene production in
vegetative tissue, seems to be ineffective in groundnut seeds. Applied
abscisic acid (ABA) maintains the dormancy of groundnut by inhibiting
ethylene production, although its effect can be competed out by cytokinin
(Ketring and Morgan, 1972). It is well established in other species that one
of the major options available to plants to prevent seed germination during
development is endogenous ABA (e.g. Kermode et ai., 1986), although
this has not been confirmed in groundnut in vivo - all Ketring and
Morgan's evidence was gained from studies with exogenously applied
ABA. Seeds with little or no dormancy can thus be expected to have put
their ethylene synthesis metabolism in place during development and also
to possess either low levels of ABA or reduced sensitivity to this plant
growth regulator.
Sharma and Sengupta (1988) have shown that a non-dormant groundnut
line possessed higher levels of ex-amylase and protease during seed devel-
opment than a dormant cultivar. While it is unlikely that this hydrolase
activity (almost certainly sequestered in the mature seed) is directly related
to dormancy control, these results are interesting in that they may rep-
resent other effects of ABA in the dormant line.
Losses of potential yield 155
TABLE 5.1 Abortion during seed development in cv. Virginia
Jumbo Runner, plants grown in adequate calcium (data reworked
from Smith, 1954)
Number of ovules Losses at each

or seeds stage (% total)
Available ovules a 4774

Fertilized ovules 4454 6.7
Ovules placed below
the soil surface 2907 25.2
Ovules resuming
development after
quiescent phase 1093 45.2
Seeds completing
development 544 11.5
a Based on the number of flowers observed, assuming two ovules per

flower.
5.6 LOSSES OF POTENTIAL YIELD
Actual seed yields in groundnut may be as low as 10% of potential yield

based on the numbers of flowers produced. Accordingly, a discussion of
seed development would be incomplete without some consideration of the
reasons for this loss. Much more detailed discussions of groundnut agro-
nomy will be presented in later chapters.
5.6.1 Losses of potential yield prior to underground pod development

As has already been described, low pollination efficiency may be a major
limiting factor for seed production in some parts of the world, but even
when most ovules become fertilized, considerable intraplant competition
means that abortion can occur throughout development. Other stages at
which potential yield was lost in Smith's (1954) study are summarized
in Table 5.1. Only 68% of flowers produced pegs, most of the rest
remaining as dormant fertilized ovules which may persist for several weeks
before withering away. However, if these ovules resume growth unusually
high seed losses occur. Pegging will then take place and development will
proceed as normal. Bunting and Elston (1980) noted that in some of the
older cultivars less than 20% of flowers may produce pegs: carpophore
elongation on the first flower of the inflorescence inhibits the development
of fruits of other fertilized flowers on the spike. Similarly, late inflor-
escences have little chance of producing successful fruit unless earlier ones
are damaged in some way. In their detailed study over several cultivars,
Bagnall and King (1991 b) showed that short days greatly increased the
proportion of pegs successfully setting pods.
In Smith's (1954) study, only one third of the pegs produced penetrated
the soil and commenced pod development (Table 5.1). Underwood et al.
(1971) showed that the penetration force of the extending carpophore was
only equivalent to 3-4 g, so that pegs have difficulty penetrating capped
soils. Normally, pegs produced by early flowers have the highest chance of
successful pod production, although if soil conditions are initially poor but
improve later during the flowering period it will be later flowers that
eventually produce successful fruit.
5.6.2 Calcium nutrition, water stress and pod development

Abortion continues throughout fruit and seed development. Many enlarg-
ing pods fail to reach maturity, while many ovules within these pods do not
set seed. Calcium nutrition in the pod region is a crucial factor in pod and
seed abortion, but calcium deficiency symptoms are often difficult to detect
above ground (Smartt, 1976).
Smith (1954) found that the proportion of empty segments in developed
pods increased from 18% to 47% under low calcium conditions. Skelton
and Shear (1971) used 45Ca++ labelling techniques to demonstrate that
calcium ions move extremely slowly to fruits developing in the soil. Under
normal soil moisture conditions there is little if any water movement to
developing fruits via the xylem and Ca++ is almost immobile in phloem.
Thus any calcium required by the developing seed has to be absorbed
directly from the soil solution by the fruit itself. Skelton and Shear found
that an increased proportion of pegs failed to set fruit on penetrating soils
deficient in calcium: 80% of all one-segment pods aborted, but all two-
segment pods succeeded in developing one seed. As with early flowers
being most successful at fruit production, this seems to be another demon-
stration of a finely tuned competitive hierarchy within the groundnut plant.
Kvi en et al., (1988) have shown that the surface area of the pod is the key
factor determining calcium uptake by developing fruits, but other factors
such as days to maturity or pod thickness are also important. Genotypes
with thick, dense hulls tend to accumulate Ca++ at the expense of the
developing seed. Lauter and Meiri (1990) have shown that the low salt
tolerance of groundnuts is probably related to the inhibition of calcium ion
uptake by developing pods via competition from Na+ for calcium exchange
sites.
Moisture around the pod zone is crucial for successful seed development
in most cultivars, but in many regions the top few centimetres of soil are
liable to dry out in the later weeks of the growing season. In a comparative
study, Wright (1989) demonstrated that water deficits in the top 8 cm of
soil caused a reduction in the seed yield of two pot-grown spanish type
cultivars even when deeper roots were adequately watered. Interestingly,
the yield components of each cultivar were affected somewhat differently:
Losses of potential yield 157
one showed a decrease in the proportion of pegs producing successful
pods, the other showed reduced yield due to a decrease in the average
number of seeds per pod. In neither case was seed weight affected. In both
cultivars, water stress in the pod zone resulted in the seeds' calcium
accumulation being reduced by 50%, as was the case in a third cultivar, a
virginia type which showed no decrease in yield as a result of a water
stressed pod zone. Wright ascribes this finding to the fact that the calcium
levels in the seeds of the third cultivar never became limiting: seeds from
these stressed plants had calcium levels comparable with the two spanish
type controls. Whether calcium is the sole limiting factor under these
conditions requires further confirmation: unexpectedly, the vegetative
growth of the virginia type cultivar was much poorer than in either of the
spanish types.
5.6.3 Plant population effects

In general, a planting population of at least 100000 plants per hectare
is recommended for groundnuts (Smartt, 1976). Many workers (e.g.
Mozingo and Coffelt, 1984) have shown the benefit of increased plant
populations, but results have been variable and clearly depend on other
factors. Kvien and Bergmark (1987) identified some of the important
parameters involved. They showed that, at 30000 plants/ha, population
density was the key limiting factor and yield improvements of nearly 30%
could be effected by increasing the population eight-fold. Plants at high
density tended to increase stem growth at the expense of assimilate parti-
tioning to reproductive tissue. At high populations, yield was sensitive to
planting date: a delay in planting by 5 weeks reduced yield by as much as
27%. At low densities plants were better able to compensate for late
sowing. Water stress reduced yield in all cases, despite the fact that plants
showed some compensation for the stress by increasing harvest index.
As previously emphasized, a key problem with the groundnut crop is the
range of pod maturities encountered at harvest. This in turn may be
reflected in changes in seed size distribution. In general, as population
density increases, numbers of seeds in the larger size grades tend to
increase (Figure 5.9). The reason for this is that high interplant compe-
tition at high densities tends to suppress the development of later repro-
ductive growth and, typically, earlier flowers are more successful at setting
seed (Kvien and Bergmark, 1987). Sung and Chen (1990) have shown that,
while cotyledon cell numbers are relatively constant, cell expansion rates
tend to be much faster in early formed pods. Nevertheless in Kvien and
Bergmark's (1987) study, between 64% and 69% of pods failed to reach
maturity in early sowings at high density, irrespective of field location.
Very immature pods will not be picked up during machine harvesting,
while slightly more advanced ones will contribute small seed to the harvest.
40
30
(/)
"t:l
Q)
Q)
(/)
20
0~
r- ':-:':':':1---
-
10 r.:':":':'
o
<7.1 7.1 7.9 8.7 9.5
Screen slot widths (mm)
Figure 5.9 The effect of plant population on seed size distribution in Florunner
peanut. Data averaged over two planting sites x two planting dates. (Calculated
from Kvien and Bergmark, 1987.)
5.7 PRODUCTION CONDITIONS AND SEED QUALITY
The next two sections of this review deal with factors affecting seed quality
in groundnut, with special emphasis on quality factors relating to planting
value. Section 5.7 reviews the effects of production conditions, while in
section 5.8 some consideration is given to the impact of storage on seed
quality.
While there is quite a large body of literature on the effects of environ-
mental conditions during development on seed yield in a range of crops,
this is less often extended to seed quality in terms of either composition or
planting value. Many of these effects may be mediated by seed size and
maturity status - for example, low soil moisture and high temperatures are
factors which may singly or in combination reduce the duration of the
reserve accumulation stage and result in smaller seeds. Alternatively,
cooler, wetter conditions may considerably delay seed maturity. While the
buffering properties of the soil may mean that some of these effects may be
less evident in groundnuts than in the cereals, it is also likely that some
important effects are overlooked owing to the indeterminate growth habit
of the crop, which results in a wide range of maturity and size classes in any
control crop grown under favourable conditions. Seed size per se may not
Production conditions and seed quality 159
always be related to maturity in physiological terms, but it is clear that
small groundnut seed have reduced food reserves and produce smaller
seedlings which often have lower survival rates. These effects may persist
until yield (Sivasubramanian and Ramakrishnan, 1974; Knauft et at.,
1990).
This section of the review attempts to collate some general ideas on this
problem under the headings of key environmental factors.
5.7.1 Temperature
Low temperatures can depress seed yield by reducing flower numbers
(Chang et at., 1985) and generally decrease rates of subsequent pod and
seed development, while high temperatures may inhibit fertilization and/or
embryo growth. Night temperatures near zero can severely impair the
planting value of windrowed groundnuts, and this loss of quality is prob-
ably caused by damage to cell and mitochondrial membranes, resulting in
increased leakage from dry seeds and higher levels of anaerobic respiration
as they begin germination (Singleton and Pattee, 1989). Again, there is an
interaction with seed maturity: small seeds respond much more severely to
chilling injury than larger ones. On the other hand, in the growing crop,
lower night temperatures may in some genotypes allow recovery from the
adverse effects of supra-optimal daytime temperatures (Smartt, 1976).
Qualitative changes in seed food reserves as a consequence of differing
temperature regimes during seed development have been noted by many
authors. Several of these effects can be explained by seed size and/or
maturity effects, as was found by McMeans et al. (1990) who showed that
sucrose concentrations in developing seeds of cv. Florunner tended to be
depressed by higher soil temperatures. As Figure 5.8 shows, increases
in sucrose levels are a relatively late event in seed development and high
temperatures are likely to curtail Stage II early. Similarly, it is not surpris-
ing that immature seeds or seeds grown under cooler conditions tend to
have lower levels of oil and protein reserves (Bovi, 1983; Nagaraj et al.,
1989). What is more interesting is that lower temperatures during seed
development can affect oil quality, favouring high proportions of poly-
unsaturated fatty acids, especially linoleic acid, at the expense of oleic acid
levels (Bovi, 1983). Another temperature related effect was noted by
Chang et al. (1985), who reported that low day-night alternations in
temperature favoured high oil yields, while higher temperature variation
favoured kernel protein contents. The physiological basis of this is not
known.
5.7.2 Water stress

It is clear from various studies that high humidity is essential for successful
groundnut seed development. This is evidenced by the fact that pods
TABLE 5.2 The effects of water stress during production on subsequent seed
quality of cv. Florigiant, (values are averages from four seasons' experiments;
adapted from Pallas et aI., 1977)
Stress conditions a Irrigation Sound mature TSW (g) Germination

water used kernels b (% ) (%)C
(cm)
~0.02 Mpa 53 66 900 92

~0.2 Mpa 46 61 940 89
~1.5 Mpa 34 55 790 82
Plants wilted
overnight 23 43 640 69 d
aWater potential at 30 em
bKernel diameter ~ 6.36 mm
cOf sound. mature kernels
dAdjusted for missing value for one season
TSW, thousand seed weight
usually develop several centimetres below the soil surface, where surface
drying effects are avoided (Smartt, 1976). Water stress will reduce ground-
nut yields but there is also good evidence that it will reduce the quality of
seeds produced. Pallas et at. (1977) demonstrated that the cultivar
Florigiant was susceptible to water potentials of -1 .5 MPa or greater.
These production conditions cause decreases in percentages of sound
mature kernels, seed weight and germinability (Table 5.2). Comparative
studies with Florunner and Tifspan showed that these smaller seeded
cultivars were less susceptible. Recent work by Ketring (1991) using other
cultivars has confirmed these results, showing that water deficits typically
result in the production of seeds with reduced early seedling growth rates.
Earlier work by this group (Ketring et at., 1978) indicated that the develop-
ment of ethylene synthesis capacity (section 5.5) was generally associated
with seed vigour and that this may be impaired by water stress conditions
during seed development.
The chemical composition of groundnut kernels may be affected by
water stress during seed production, but there may also be major inter-
actions between genotype and the timing of the water deficit. Ross and
Kvien (1989) looked at changes in sugars and phenolics in kernels sub-
jected to water stress during the growth of the parent plants. Typically,
mid-season drought caused increases in fructose, glucose and phenolic
compounds in mature seeds, but the effects on sucrose contents were
highly variable. Four out of eight cultivars evaluated showed increased
kernel sucrose contents as a result of water stress at some stage of develop-
ment. One line showed significant decreases in seed sucrose levels when
water-stressed 50-80 days after sowing, while the cultivar Tifton 8 showed
wide variations in sucrose contents, as compared with seeds from
Production conditions and seed quality 161
unstressed controls, with around 40% increases when stressed at 20-50
days, but a decrease of equal magnitude when parent plants were stressed
at 50-80 days.
5.7.3 Mineral nutrition

Specific effects of various mineral element deficiencies during production
on subsequent seed quality have been documented by several authors. The
importance of adequate calcium nutrition in preventing abortion has
already been mentioned, although adequate Ca++ may also be important
to maintain the planting value of set seed. Cox et at. (1976) found that a
seed calcium content of 420 ppm or higher was necessary for high germina-
bility in cv. Florigiant. Ca++ concentrations below this value resulted in
decreased germination, falling to almost half the control value when the
calcium content was reduced to 200 ppm. Maeda et at. (1986) noted that
liming caused a reduction in seed size in ground nuts produced in Brazil
(presumably due to pH effects, cf. Smartt, 1976) but that these seeds had
improved germinability and vigour.
Interactions between calcium and boron deficiency in groundnut are well
known. Cox and Reid (1964) and Harris and Brolmann (1966) showed that
blackening of plumules was caused by calcium deficiency whilst cotyledon-
ary damage, especially hollow heart, was largely a function of boron
deficiency, although lack of Ca++ may also contribute to discolouration of
the cotyledons (Table 5.3). Harris and Brolmann (1966) demonstrated that
plumule damage due to Ca++ deficiency was caused by disruption of
vascular tissue at the base of the plumule.
TABLE 5.3 Influence of calcium and boron nutrition during seed development on
embryo damage in groundnuts (data from Cox and Reid, 1964)
Damaged kernels (%)
Site of damage 0.029% Ca++ 0.039% Ca++
8ppmB 12 ppm B 8ppmB 12 ppm B

Plumules 34 30 9 7
Cotyledons 14 8 6 1
Boron deficiency mayor may not affect seed yield, depending on cultivar
and other cultural factors (Rerkasem et at., 1988). For instance, there may
be significant interactions with other mineral elements besides calcium.
Patil et at. (1987) reported that combined applications of boron and
phosphate were much more effective than either element alone in improv-
ing seed numbers, seed weights, protein and oil contents from groundnuts
grown in boron deficient soils in India.
5.7.4 Other management factors
Plant spacing has been shown to affect the maturation pattern of some
cultivars of groundnut. For example, Mozingo and Coffelt (1984) found
that increased interplant competition within rows resulted in plants setting
fewer late pods, which allowed an increased proportion of large kernels
without an overall reduction in yield. However, extensions of this kind of
work into attempts to manipulate ground nut seed yield or quality via
chemical manipulation, e.g. the anti-auxin Kylar (Wynne et ai., 1974),
seem to have been largely unsuccessful. Worthington and Smith (1974)
have shown that foliar applications of chemicals to the parent crop can
affect the nutritional value of groundnut kernels: Kylar tended to decrease
the proportion of linoleic acid, while the fungicide benomyl had the
opposite effect.
5.8 QUALITY FACTORS AND THE PROCESS OF SEED

DETERIORATION
5.8.1 General principles of groundnut storage

It is well known that groundnut seeds are more difficult to store than most
other seed crops, with the exception, perhaps, of soybean (Barton, 1961).
Any losses in seed viability will be preceded by considerable losses in
planting value at earlier stages of storage. This poor storability is attributed
to the high oil content of these seeds (e.g. Gelmond, 1971b) and many
studies have focused on the relationship between oil composition and
storability of groundnut kernels and of groundnut oil itself. This topic will
be discussed in section 5.8.3, but it is important to preface it with a short
general discussion on the principles of seed storage and then to review
present ideas about the mechanisms involved in seed deterioration.
The basic principles of seed storage are well known. Essentially, the key
points are to process seed carefully, clean it well and then keep seed
moisture content and storage temperature as low as possible. In standard
commercial practice, target seed moisture contents (SMCs) are around 8%
for unshelled seeds or 6% for shelled kernels, which are usually more
difficult to store (Woodroof, 1973; Shewfelt and Young, 1977; Navarro et
ai., 1989). Shelled kernels are normally traded in the US food industry at
an SMC of around 7.5% (Woodroof, 1973) but seeds intended for planting
should be handled more carefully. It should also be recognized that relative
humidity (RH), rather than seed moisture content per se, is the key
criterion for many physiological events and microfloral associations during
seed storage. Groundnut seeds, having such a high oil content, equilibrate
at much lower SMCs for a given RH than do cereal grains (Halloin, 1986).
Standard practice in the US is to store the crop in shells under controlled
Quality factors and seed deterioration 163
RH in cool stores. If financial constraints on storage facilities prevail, more
attention should be given to keeping seed moisture and store RH low than
reducing the storage temperature.
Several empirical studies have been carried out on the effects on seed
germ inability of different storage conditions. In comparing different
accounts in the literature, it should be remembered that there may be
differences in the storage behaviour of different cuitivars, the quality of the
particular seed lot studied, the method of evaluation of germinability
subsequent to storage and/or possible contamination of the lot by storage
microftora. Recent work includes that by Navarro et al. (1989), who
suggest that seed moisture content should not exceed 8% if shelled seeds
are to be maintained at 90% germination for 6 months at 15°C, but that a
1% reduction in seed moisture will more than compensate for a 10 °c rise
in storage temperature. In earlier studies, Gelmond (1971b) had shown
that seeds stored at 4°C and 30% RH (expected seed moisture content
=4.5%) still retained good planting value after three years' storage. Going
back further in the literature, Barton (1961) recommend storage SMCs as
low as 3% when storage temperatures exceed 26°C: however, reducing the
SMC of unshelled kernels below 6% makes seed very prone to mechanical
damage, especially loss of the thin testa on subsequent processing (Gel-
mond, 1971b).
Vertucci and Roos (1990) attempted to locate applied research of this
type within an appropriate theoretical framework based on the physiologi-
cal status of the seed in store. They calculated that, at an SMC of around
5%, the viscosity of water within the peanut kernel should be so high as
effectively to inhibit any deleterious chemical reactions in an aqueous
system. In the same study, they showed that reducing SMCs below 4%
caused the rate of deterioration of groundnut kernels to begin to increase
again, presumably due to the loss of bound water which is essential to the
structural integrity of major macromolecules. Accordingly, it can be con-
cluded that a moisture content of 4-5% is the ideal safe level for shelled
kernels but, as indicated above, maintaining SMCs as low as this may be
inappropriate for short-term bulk seed storage, especially as kernels are
liable to skin at this level. However, such information is invaluable for the
practice of the long-term seed storage required for genetic conservation.
5.8.2 Mechanisms of deterioration in stored groundnut

Seed deterioration is a complex matrix of processes involving both oxidat-
ive and hydrolytic damage. While many facets of cellular structure and
metabolism may be affected by the deterioration process, key events seem
to be either loss of membrane integrity or genetic damage. One fundamen-
tal problem in the study of the physiology of deterioration is the difficulty
of separating secondary deteriorative events (the consequences of earlier
damage) from primary damage-initiating reactions. The relationship
between these events may vary according to cultivar, the prestorage history
of the seed lots and, in particular, the nature of the storage conditions
themselves (e.g. Priestley, 1986). Sometimes, too, interpretation of experi-
mental data may not be as straightforward as it appears. For example,
observations of increased leachate from deteriorating peanuts during early
imbibition (e.g. Nautiyal et al., 1988) are usually interpreted to be an
indication of membrane deterioration, but they may equally be due to
increases in solutes available to leak out of rehydrating/reorganizing cells.
Pattee et al. (1981), for instance, reported that sucrose and stachyose levels
increased dramatically in stored immature seed but not in mature kernels.
Further complications arise when it is recognized that seeds are capable of
some degree of self-repair at higher moisture contents and also have a
considerable array of detoxification and antioxidant systems.
Vertucci and Roos (1990) have demonstrated that oxidative phosphoryl-
ation activity becomes negligible in groundnut at =16% SMC. It is unrea-
sonable, therefore, to assume that much repair activity will occur in
groundnuts at moisture contents below this level. However, it is essential
to store them at much lower moisture contents than this because of high
rates of physiological deterioration and the proliferation of seed storage
fungi, especially Aspergillus and Penicillium species. There may be signifi-
cant growth of Aspergillus when the SMC of groundnut is as low as 9%
(Halloin, 1986). Although tannins present in the testa may inhibit the
germination of fungal spores, and other phenolic leachates also have
fungistatic properties (Sanders and Mixon, 1978; Kalaichelvan and
Mahadevan, 1987), these defence mechanisms are of limited value in
prolonged storage at raised seed moistures and their effectiveness may vary
considerably with cultivar and the condition of the seedlot. The basis of
good seed storage, therefore, is to attempt to slow down the various
deteriorative events by keeping moisture contents as low as possible.
Reducing storage temperature will, of course, have similar effects.
It is generally accepted that polyunsaturated fatty acids are more prone
to oxidative damage than other stored lipids within seeds (e.g. Harman and
Mattick, 1976; Bewley, 1986). In the presence of oxygen, free radicals
(highly reactive molecular groups with an unpaired electron) can attack
fatty acid chains containing more than one double bond to form lipid
peroxides in a potentially devastating chain reaction sequence (Wilson and
McDonald, 1986; Benson, 1990). Both storage and membrane lipids may
be affected in this way, with damage to cell membranes being particularly
critical in seed deterioration, especially as lipid peroxides may also attack
membrane proteins. Lipid peroxidation may occur spontaneously, initiated
by the action of light or metal ions (autoxidation), or it may be enzymically
driven by the lipoxygenase enzyme. There is considerable controversy,
however, on the relative importance of free radical damage in seed deterio-
ration compared with other types of damage (e.g. Wilson and McDonald,
1986). For instance, free radicals are likely to be quenched by free water in
Quality factors and seed deterioration 165
the cell, while natural antioxidants and detoxification systems are also
present.
Based on observations and research done by the food industry, it is often
assumed that autoxidation is a key deteriorative event in the dry storage of
intact groundnut kernels as well as in their products. However, very few
biochemical evaluations of deterioration mechanisms in kernels appear to
have been published and these seem inconsistent with each other. From
the composition of volatiles released from kernels stored at 6% SMC and
ambient temperatures, Pattee et al. (1971) concluded that autoxidation
reactions were of minor importance and that both lipoxygenase and other
enzyme activities were necessary to produce the range of compounds
detected. While they legitimately argued that, even at 6% SMC, there may
be localized areas of significantly higher moisture content to permit activity
of these enzymes, it is not possible to demonstrate the in vivo activity of
these enzymes unequivocally.
Pearce and Abdel Samad (1980) approached the problem by looking at
substrate depletion in deteriorating groundnut stored under either good or
adverse conditions. In no case were they able to demonstrate any evidence
of lipid peroxidation damage, despite the fact that there was considerable
depletion of phospholipid and loss of membrane integrity - events which
preceded losses of germinability. They concluded that hydrolytic damage,
rather than autoxidation or lipoxygenase-driven reactions, were the most
important events in deterioration. In contrast, Chen and Fu (1986)
reported that ageing both increased lipid peroxide levels and caused a
decrease in natural antioxidants (e.g. glutathione, ascorbate and super-
oxide dismutase). Of particular interest is their finding that treatment of
kernels with ascorbate or glutathione solutions improved seed vigour.
Clearly we need to know much more about the specific mechanisms of
deterioration in groundnut seeds.
5.8.3 Oil content and storability of groundnuts

Despite the uncertainties outlined in the previous section, it does appear
that there is a correlation between polyunsaturated fatty acid levels and
storability of groundnuts. As most groundnut cultivars have only trace
amounts of polyunsaturated fatty acids other than linoleic acid (section
5.4), levels of the latter compound in relation to the other major fatty acid
component, oleic acid, are often used as an index of storability, viz the
C 18 : 1: C 18 :2 ratio or oil stability index. Low values of this index are generally
associated with reduced storability of both intact kernels and groundnut
products (Shewfelt and Young, 1977).
In general the oil stability index seems much higher for virginia runner
types than for virginia bunch, valencia or spanish peanuts (Raheja et al.,
1987; Nagaraj et al., 1989). Table 5.4 shows the data range obtained by the
former group of authors in their comprehensive study. Moore and Knauft
TABLE 5.4 The range of oil contents and stability indices of three types of
groundnuts determined in a survey of 82 cultivars (data from Raheja et a\., 1987)
Virginia Virginia Spanish

runner bunch bunch
Oleic acid
(as % total oil content)a 50.6-54.7 37.6-49.1 36.3-49.7
Linoleic acid
(as % total oil content) 29.7-33.4 33.8-46.7 30.0-44.3
Oil stability index b 1.53-1.84 0.82-1.45 0.83-1.53
Iodine values c 99.2-100.9 101.5-114.2 93.7-111.2
aTotal oil contents of the three types were very similar and around 50% dry weight
bOil stability index is the C]S,]:C]8,2 ratio
clodine valucs are an indication of the proportion of double bonds in the oil or fat sample, but
are also affected by the proportions of free fatty acids present (Cocks and van Rede, 1966)
(1989) have identified two recessive genes in ground nut which indepen-
dently influence oleic acid levels in kernels. The expression of either gene
can raise the proportion of oleic acid in peanut oil to around 70% , which is
likely to improve the storability of both extracted oil and intact kernels but
impairs the nutritional quality of the product.
Apart from genetic variation, factors during seed production can also
affect oil stability. Young et al. (1972) noted that mature kernels tended to
be higher in 18:0 and 18: 1, but lower in 18:2 fatty acids. Similarly, Mozingo
et al. (1988) reported that larger kernels tended to have higher C18:1:C18:2
ratios. This group was also able to show that growing season and field
location both affected this parameter, but relevant climatic data were not
supplied. Bovi (1983), however, observed that the C18:1:C18:2 ratio was
reduced in kernels growing in cooler locations.
As might be concluded from the previous sections, it is a dangerous
assumption to infer that oil stability is entirely a function of C 18 : 1 :C18 :2
ratios. In a major study, Worthington et al. (1972) examined the corre-
lation between the time taken for expressed oil to become rancid (the
autoxidation induction period) and the lipid composition of that oil. Late
maturing types were generally higher in oleic and lower in linoleic acid
levels, but it can be seen from some of their data (shown in Table 5.5) that
the importance of oleic:linoleic acid ratios varies considerably with grow-
ing season. In 1967 and 1968 it accounted for a high proportion of the
variation in oil stability, but in 1965 correlations were very poor. Clearly
other factors are involved - no doubt levels of natural antioxidants are
important, but what else is involved and the precise nature of interactions
with environmental conditions during seed production are still a matter of
debate, as is the exact relationship between this type of data and the
situation in whole kernels.
References 167
TABLE 5.5 Mean oleic and linoleic acid compositions over 82 groundnut geno-
types in three different production years and the correlation between the autoxidation
induction period (AlP) and the CI8:/:CI8:2 oil stability index. (Standard errors of
individual means are shown in brackets. Data from Worthington et aI., 1972)
Growing season
1965 1967 1968
Oleic acid
(% total oil content) 50.1 (± 10.5) 49.1 (± 9.03) 50.3 (± 10.09)
Linoleic acid
(% total oil content) 24.5 (± 8.20) 30.1 (± 7.21) 30.4 (± 8.33)
Correlation between AlP
and C 18 : I :C I8 :2 ratio
(R2 value) 10% 73% 53%
R2, coefficient of determination
5.9 SOME FINAL CONCLUSIONS
From the foregoing discussion, it is clear that we have a reasonably

comprehensive picture of the morphological and compositional changes
occurring during seed development in groundnuts. We are also beginning
to build up a body of knowledge on how to manipulate oil content and
storability by a combination of breedin!; programmes and appropriate
agronomy. There are, however, two key areas where there are major gaps
in our knowledge. Firstly, we know almost nothing about how seed devel-
opment in groundnut is controlled. We know very little about the role
played by endogenous plant growth regulators in the reproductive biology
of this species and virtually nothing about how they mediate intraplant
competition and losses of potential yield. This is certainly an area ripe for
future research. The second area for future work is to improve our
understanding of how pre- and post-harvest environmental factors affect
seed quality, particularly in terms of planting value. In this instance there is
scope for both careful empirical research and some basic physiological
studies, especially on seed deterioration in store.
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CHAPTER 6
The composition and nutritive value

of groundnut kernels
G.P. Savage and J.I. Keenan
6.1 INTRODUCTION
Groundnuts were originally considered to be food for animals; then they

were used as food for slaves. They have now become an important source
of protein in many developing countries. A summary of the important food
uses of groundnuts and the fortified foods that use groundnuts or ground-
nut protein has been given by Singh and Singh (1991).
The use of groundnuts by the US armed forces as peanut butter and as
snacks has boosted their popularity in western countries as well. It should
be remembered that groundnuts are primarily grown as a source of edible
vegetable oil; they are the fourth most important source of edible oil and
the third most important source of vegetable protein.
6.2 PROXIMATE COMPOSITION
The proximate composition of groundnut kernels and a number of

products made from them are presented in Table 6.1, which contains
a summary of the accepted values that can be found in the literature.
(Where significant variations from these values occur, they will be dis-
cussed in the appropriate section.) The type of processing alters the
gross composition significantly, from full fat flour which contains
essentially the same amounts of the major classes of nutrients as the
raw kernels, to protein isolates which contain more than 90% crude
protein (N * 6.25) on a dry weight basis as well as traces of other com-
ponents.
The Ground-nut Crop: A scientific basis for improvement. Edited by 1. Smartt. Published in
1994 by Chapman & Hall, London. ISBN 0412408201.
174 The composition and nutritive value of groundnut kernels
TABLE 6.1 Proximate composition of groundnuts and groundnut products
(Natarajan, 1980)
Kernels Full fat Defatted Protein Protein

flour flour concentrate isolate
Moisture 5.0 2.5 7.5 4.5 3.4

Protein 30.0 27.4 57.0 60.0 95.0
Fat 48.0 44.4 0.6 8.0 0.5
Carbohydrates 15.5 21.5 30.0 18.5
Crude fibre 3.0 2.3 4.6 4.0
Ash 2.0 2.0 4.6 2.1 0.5
6.3 OIL AND FATTY ACIDS
Groundnuts are valued for their high quality oil content. About two thirds
of the world production of groundnuts is utilized as an edible oil, making it
one of the world's leading oil seed crops. Most of the oil is found in the
cotyledons which comprise approximately 72.4% of the kernel (Fedeli et
at., 1968; Woodroof, 1983). Groundnuts are also consumed whole as
snacks, and in confectioneries and peanut butter. In this form, they are
usually roasted before use. This accentuates the flavour which contributes
to consumer acceptance. Studies have shown that the oil content and
composition undergo very little, if any, change as a result of roasting
(Hoffpauir, 1953; Iverson et at., 1963; Sekhon et at., 1970; Khalil and
Chughtai, 1983).
6.3.1 Quantitative changes in oil content and fatty acid composition

Studies have shown considerable compositional and quantitative changes
in the lipids and fatty acids of groundnut seeds during the growth period to
maturity, i.e., 15-80 days. The oil percentage increases directly in pro-
portion to the increase in dry seed weight, which is most rapid between
7 and 8 weeks, reaching a maximum at 11 weeks (Sanders, 1980a). From 7
weeks onward, lipid synthesis replaces starch as the dominant reserve
accumulation mechanism (Pickett, 1950; Abdel Rahman, 1982b). The
maturity of the seed also has an important effect on the composition of the
oil (Pickett, 1950). The free fatty acid levels decrease (from approximately
4.5% to 0.7%), offset by triglyceride formation. The oleic acid (18:1)
levels of the triglycerides and compound lipids increase rapidly. Shibahara
et at. (1977) report a corresponding decrease in the proportions of linoleic
(18:2) and palmitic acid (16:0). Abdel Rahman (1982a,b) and Sanders
(1980b) found the linoleic acid (18:2) level to be changeable. The greatest
change apart from oleic acid was in the levels of behenic acid (Sanders,
1980b).
Oil and fatty acids 175
These changes can be important, as oil stock ground nuts are usually
those that are rejected or diverted from edible channels. Rejection can be
for a number of reasons, including small kernel size (often an indication
of kernel immaturity). Weathered or insect infected nuts (a sign of
inadequate storage) are also used (Woodroof, 1969). Whiston et at. (1959)
showed decreased oil content and increased free fatty acid levels in kernels
(especially those that were damaged) stored for 24 months.
6.3.2 Analysis of kernel content

Given the changing nature of the fatty acids in immature kernels, It IS
essential to select mature kernels for analysis in order to obtain highly
reproducible results. Typically they are smooth, with little or no wrinkling
of the testa surface. They must also have a dark-coloured interior pericarp
surface with a thin, faded testa (Worthington et at., 1972; Young and
Waller, 1972). The internal distribution of the oil varies and this can be
significant when only part of the kernel is to be sampled. The tip opposite
the germ contains the most unsaturated oil; the centre of the kernel has
been reported as having the lowest iodine value, indicative of more satu-
rated fatty acids (Young and Waller, 1972). Ideally one quarter to one half
of the kernel is taken from the end opposite to the germ end.
Oil can be extracted by mechanical or solvent means from the kernel.
A silver-plated Carver press can be used to expel the oil. Alternatively,
solvent extraction (with petroleum ether or chloroform-methanol) can be
carried out in a Soxhlet apparatus for 18 hours. Chloroform-methanol
extraction always extracts the maximum amount of lipid (Varma and
Kumar, 1968; Sanders, 1980). A comparative study by Wallerstein et at.
(1989) did not show any significant differences between Soxhlet extraction
and the modified Gerber method, which involves sulphuric acid digestion
of the seed pulp and phase separation of the oil. Other methods include a
rapid butyrometric method (Shukla et at., 1980) or oil analysis by nuclear
magnetic resonance, a non-destructive and rapid analytical method which
would be of use in breeding programmes (Jambunathan et al., 1985).
Individual fatty acid content is determined after the preparation of
methyl esters from the oil samples by transmethylation with 3% sulphuric
acid in methanol. Analysis is then by gas liquid chromatography (Worth-
ington and Holley, 1967).
6.3.3 Total oil content

Groundnuts can be divided into three distinctive types on the basis of plant
characteristics and habits of growth: spanish (bunch), virginia (spreading
bunch) and runner. Data from several sources (Hoffpauir, 1953; Hopkins
and Chisholm, 1953; Iverson et al., 1963; Sekhon et al., 1972a; Sekhon et
al., 1973; Woodroof, 1983; Taira, 1985; Wallerstein et ai., 1989) show that
the oil content of these three varieties differs to some extent, as follows:
Spanish 42.0-53.8% of dry weight
Virginia 45.0-58.6% of dry weight
Runner 41.23-53.6% of dry weight
In a study covering eight countries, Taira (1985) reported a significant
geographical effect on the oil content of a spanish cultivar which ranged
from 46.9% to 58.6%.
Other authors have reported oil content without specifying the ground-
nut type. These range from 42.3-56.5% (Crawford and Hilditch, 1950;
Iverson et al., 1963; Rao et al., 1965; Oke, 1967; Sekhon et al., 1970;
Sekhon et al., 1972b; Badami et al., 1980; Abdel Rahman, 1982a; Khalil
and Chughtai, 1983) while Roberson et al. (1966) reported 44.4-54.4% oil
content in 30 commercial peanut butter samples.
Because two thirds of the groundnuts cultivated worldwide are grown for
their oil content, a level of more than 50% oil is desirable. Plant breeding
techniques are now being used to improve the quantity (and quality) of oil
in cultivars.
6.3.4. Oil composition

The quality of the oil IS Important from the point of view of human
nutrition as well as the stability of the oil during storage.
(a) Saponifiable fraction

The saponifiable fraction of groundnut oil consists mainly of triglycerides
of long-chain fatty acids. The phospholipids lecithin and cephalin also
occur but are thought to be less extractable from cured seed (Sanders,
1980b). They are also thought to settle out of crude oil. Alkaline hydrolysis
of oil or fat gives a 'saponification number' which indicates the nature of
the fatty acids present, since those with longer carbon chains liberate less
acid for each gram of oil hydrolysed. The saponification value is only of
interest if the groundnut oil is going to be used for industrial purposes as it
has no nutritional significance. Saponification numbers reported for
ground nut oil range from 190 (spanish type) to 298.8 (Crawford and
Hilditch, 1950; Hopkins and Chisholm, 1953; Sekhon et al., 1970).
Fatty acid composition

Eight fatty acids account for more than 98% of the total fatty acid compo-
sition of groundnut oil (Worthington et al., 1972; Woodroof, 1983).
Palmitic, oleic and linoleic account for more than 80% of total fatty acids
after 30 days growth (Shibahara et al., 1977). They eventually contribute
approximately 90% of total fatty acids at kernel maturity (Young and
Waller, 1972; Sekhon et al., 1972a,b) but varying ratios of these three fatty
acids are found within the three major types (Table 6.2). The virginia type,
which is later maturing, and the runner type are generally higher in
monounsaturated fatty acids (oleic acid). The earlier maturing spanish type
has a higher linoleic acid (i.e. polyunsaturated) content but total saturated
fatty acids are also higher (Woodroof, 1983; Worthington et at., 1972).
Virginia cultivars have been reported to contain higher levels of linoleic
than oleic acid (Sekhon et at., 1973), which is not observed in spanish
cultivars (Sekhon et at., 1972a).
Strains grown under similar climatic and agronomic conditions still show
varietal differences which have been attributed to genetic causes (Sekhon
et at., 1972b). This makes manipulation of the oil content and composition
through plant breeding techniques a practical objective (Sekhon et at.,
1972a). An alternative has been the development of mutant breeding using
gamma-irradiation (35-75 kR dose range) (Sharma et at., 1981). While
having no effect on the overall oil content, the process produced mutants
with a higher oleic and lower linoleic acid content than the parent strain.
The two mutants showed a substantial increase in oleic acid, with a
corresponding decrease in palmitic acid, a ratio which is nutritionally more
desirable. The other fatty acids showed minor variation.
In a study of 30 commercial peanut butters, Roberson et at. (1966) found
comparable composition and fatty acid levels, with the small differences
between various brands probably due to the variety of groundnuts used or
the amount of added hydrogenated fat. Little difference was found be-
tween smooth, medium and chunky blends of the butters.
Differences in fatty acid composition

It has been reported that the composition of groundnut oil is influenced by
environmental and seasonal effects in addition to varietal differences,
genotypic variation and maturity (Young et at., 1974c; de Bertorelli, 1976;
TABLE 6.2 Fatty acid composition of groundnut oil (% of total fatty acid)
Fatty acid type Spanish Virginia Runner Unspecified
16:0 8.2-18.4 6.0-13.S 8.6-13.3 6.0-20.12

16:1w9 0.1-2.3 0-1.1 0.1--0.2 0.1-1.47
18:0 1.1-6.2 1.0-4.9 2.0-3.0 0.8-S.3
18:1w9 36.4-S3.1 36.7-61.1 40.9--60.4 3S.7-71.S
18:2w6 20.S-40.3 21.6-48.3 21.0-3S.2 13.0-41.1
18:3w6 0.1--0.3 O.l--O.S 0.1--0.2 O-1.S
20:0 1.0-2.1 1.1-1.8 1.0-1.5 0.9-3.S
20:1w9 0.7-1.4 1.1-1.8 0.9-1.9 0.6-2.0
22:0 3.0-4.4 2.7-3.7 2.8-3.9 1.3-S.1
24:0 0.9-2.0 1.5-2.0 1.2-2.6 0.6-S.9
Based on data from Hoffpauir, 1953; Hopkins and Chisholm, 1953; Iverson et al., 1963;
Worthington and Holley, 1967; Sekhon et al., 1972a; Sekhon et al., 1972b; Worthington et al.,
1972; Sekhon et al., 1973; de Bertorelli, 1976; Badami et aI., 1980; Kritchevsky et al., 1981; Abdcl
Rahman, 1982a; Nouvelot et al., 1983; Woodroof, 1983; Taira,. 1985; Wallerstein et al., 1989.
Taira, 1985). These effects include annual climatic fluctuations, variable
soil conditions and agricultural practices. The environment (especially
temperature) during seed formation can modify both the oil content and
the fatty acid distribution pattern in a given cultivar (Worthington et at.,
1972; Liu et at., 1984). Taira (1985) showed that the daily mean tempera-
ture during ripening correlated negatively with the linoleic acid content.
Young et al. (l974c) reported several significant first and second order
interactions with the three major fatty acids involving variety, location and,
most noticeably, soil moisture conditions. Inanga et at. (1990) studied the
effect of calcium levels in the soil during the growing period. They found
that, while calcium deficiency can lower phospholipid content, it has no
effect on the fatty acid composition of the triglyceride.
Linolenic acid
One characteristic of groundnut oil is its low linolenic acid (18:3) content.
Studies have shown that authentic groundnut oil contains considerably less
than 1% linolenic acid (0.02-0.3%) and that this value is not greatly
influenced by geographical distribution (Worthington and Holley, 1967;
Worthington et at., 1972; Worthington, 1977; Taira, 1985). In such small
amounts it is able to exert an effect on the oil flavour without contributing
to oxidative rancidity. Ideally it can also be used for the identification of
authentic groundnut oil and the detection of adulteration. However, the
acceptance of a value of less than 1.0% linolenic acid would still permit
the adulteration of groundnut oil with 10% or more soya bean oil as the
linolenic acid content of the mixture would still be less than 1% (Worth-
ington, 1977).
Several authors do not report linolenic acid levels, while others such as
Iverson et af. (1963) were unable to detect it. The methodology used to
detect the fatty acids is important. Ideally GLC (gas liquid chromatogra-
phy) liquid phases with polarity characteristics that allow elution of C18
fatty acid methyl esters prior to the elution of the C20 series should be
used; otherwise there may be difficulties in distinguishing between linole-
nic acid, eicosanoic acid (20: 1) and, less frequently, arachidonic acid
(20:0). Hoffpauir (1953) reported an arachidonic acid range of 2.4-4.0%.
The usually reported range is 0.9-3.5%. This would suggest misidentifica-
tion of linolenic and eicosanoic acids as arachidonic acid. Similarly,
Badami et af. (1980) reported that one cultivar contained 1.46% linolenic
acid but made no mention of eicosanoic or behenic acids.
(b) U nsaponifiable fraction

The unsaponifiable fraction of groundnut oil consists of a mixture of classes
of compounds including hydrocarbons, tocopherols, sterol esters, free
sterols, glycolipids and others. This fraction has been reported as ranging
from 0.7% to 1.6% (Crawford and Hilditch, 1950; Hoffpauir, 1953;
Hopkins and Chisholm, 1953; Fedeli et at., 1968; Badami et at., 1980),
which indicates that individual compounds within a class are usually pres-
ent in only trace amounts.
Phytosterols account for 0.16-0.25% of the total oil content and are the
main component of this fraction (Hoffpauir, 1953; Worthington and
Hitchcock, 1984; Kritchevsky et at., 1981). The main components of the
free sterol fraction are f3-sitosterol, campesterol and stigmasterol. Steryl
esters also account for a portion of this fraction (Fedeli et at., 1968). The
characteristic odour and flavour of ground nut oil is attributed to the
presence of extremely minute quantities (1.8 g/ton) of higher hydrocarbons
(C 1s H 30 and C 19 H 3S ). Squalene is the major component of this fraction
(Fedeli et al., 1968; Worthington and Hitchcock, 1984). The tocopherols in
this fraction have an important role in oil stability (discussed later in this
chapter). They have another important role, shown by their effect on other
vitamins - mainly vitamin A.
6.3.5. Oil quality

Vegetable oils are promoted as being healthy on the basis of their varying
degrees of unsaturation. This can be measured by the number of grams of
iodine absorbed across the double bonds by 100 g of oil. Reported iodine
values for groundnut oil range from 84.5 to 103.0 (Crawford and Hilditch,
1950; Hoffpauir, 1953; Hopkins and Chisholm, 1953; Fedeli et at., 1968;
Sekhon et at., 1970; Worthington et al., 1972; Khattab et at., 1974; Badami
et al., 1980). The iodine value only gives an indication of the total amount
of unsaturated fatty acids in an oil. It is more important to determine the
amounts and ratios of the predominant fatty acids.
A number of tests can be used to determine the quality and freshness of
groundnut oil. The oil from mature, freshly ripened groundnuts should
contain less than 1% free fatty acids. Therefore the presence of higher
levels in fresh oil may be an indication of tissue damage and the subsequent
degradation of lipids by lipolytic enzymes. This usually occurs during
storage (Crawford and Hilditch, 1950; Sanders, 1980a). The free fatty acid
content of groundnut oil (as % oleic acid) has been reported to range from
0.1 % to 7.2% (Crawford and Hilditch, 1950; Hoffpauir, 1953; Rao et al.,
1965a; Sekhon et al., 1970; Sanders, 1980; Kritchevsky et at., 1981).
Phillips and Singleton (1978) noted a problem with the hydrolysis of fatty
acid triglycerides when measuring free fatty acids, which could account for
the higher values. Their method avoided this problem. Similarly, when
exposed to air, triglycerides containing highly unsaturated fatty acids tend
to undergo a complex process of autoxidation, resulting in a number of
products that are responsible for the off-taste characteristic of rancidity.
Fresh oils should have a negligible peroxide value, e.g. 0.68 mmollkg
(Khattab et al., 1974).
Whether oil is stored on the shelf for long periods or submitted to
accelerated oxidation tests by exposure to 100°C in air, an increase in free
fatty acid content and peroxide value will occur (Varma and Kumar, 1968;
Arya et al., 1972; Wallerstein et al., 1989).
The degree of un saturation of groundnut oil is due almost entirely to the
oleic and linoleic acid contents and may be expressed as a ratio of these
acids, i.e. oleic/linoleic (or 18: 1/18:2) ratio. These two fatty acids can
account for up to 80% of the total fatty acids (Young and Waller, 1972;
Abdel Rahman, 1982a; Woodroof, 1983). A high linoleic acid content,
with its two unsaturated bonds, is undesirable because it decreases the
shelf life of groundnut products (Sekhon et al., 1972a,b). Therefore a
higher ratio of oleic to linoleic acid in groundnut oil and other groundnut
products is considered an indicator of a more stable product. However, in
nutritional terms, a higher linoleic acid content is more desirable because
of its hypocholesterolaemic effect (Young and Waller, 1972; Taira, 1985).
Oleic/linoleic ratios, calculated from the literature, range from 0.76 to 5.5
(Hoffpauir, 1953; Worthington and Holley, 1967; Sekhon et al., 1972a;
Sekhon et al., 1972b; Sekhon et al., 1973; Iverson et ai., 1963; Badami et
ai., 1980; Kritchevsky et ai., 1981; Abdel Rahman, 1982a; Taira, 1985;
Wallerstein et ai., 1989).
Plant breeding can produce a groundnut oil with a desirable level of
unsaturation (Sekhon et ai., 1973). A rapid microanalytical method for
determining the oleic/linoleic ratio by GLC has been developed. It uses
only a small portion of kernel, which means the remainder can be planted
for reproduction (Young and Waller, 1972). However, Worthington et ai.
(1972) conducted a study which indicated that yearly or seasonal variations
in environmental conditions also have an unknown but very pronounced
effect on oil stability, largely unrelated to the levels of linoleic acid in the
seed.
The tocopherols, which are powerful antioxidants, also contribute to the
stability of groundnut oil and other groundnut products containing oil. It is
possible that antioxidants and/or synergists other than tocopherols are
present in the crude oil and that these compounds are removed in refining
(Worthington et ai., 1972). Shibahara et ai. (1977) found the amounts of
tocopherols increased in direct correlation with the accumulation of un-
saturated fatty acids of triglycerides in the seeds from 40 days to maturity.
They normally range from 0.3 to 0.5 g/kg (Hoffpauir, 1953). No
j3-tocopherols are found in groundnut oil but a-0.18 to 0.3 g/kg, )'-0.18 to
0.22 g/kg and o-tocopherols have been identified The a-form is found
predominantly at the early stage of seed development, whereas the ),-form
is found at maturity.
The degradative changes that take place in the un saponifiable fraction
after oxidation can inhibit the activity of pancreatic lipase on the oil. The
antioxidant properties and sterol content decrease markedly in groundnut
oil (Ramamurti and Banerjee, 1950) on storage. This could contribute to
the depressed growth seen in experimental animals fed rancid groundnut
oil.
Another factor involved in oil quality is related to its use as a cooking
medium. Refined groundnut oil contains mostly monomeric material, no
dimers and only a very small quantity of polymers. As the oil is heated up
to 260°C, the monomers decrease, the polymers increase and the dimers
appear in increasing amounts with temperature and time. Similarly the
percentage decrease in the rate of in vitro digestibility falls to 52% at
260°C, attributable to the formation of compounds which inhibit lipolysis.
Peroxide values and free fatty acid values increase (Khattab et al., 1974)
while linoleic acid is reported to decrease by as much as 95% after 6 hours
heating at 260°C. Iodine values also decrease. Different values would
suggest the quantity of oil and surface area in contact with the air during
heating are relevant to the degree of deterioration (Arya et al., 1972;
Vidyasagar et al., 1974; Khattab et al., 1974). In contrast, Kokatnur et al.
(1964) reported a negligible increase in peroxide value after heating
groundnut oil for 8 hours at 200°C.
6.3.6 Groundnut oil and human health

Groundnut oil, despite its unsaturated fatty acid components, has been
found to be extremely atherogenic to rhesus monkeys (Vesselinovitch et
al., 1974), rabbits (Kritchevsky et al., 1971 Kritchevsky et al., 1973;
Kritchevsky et al., 1984) and rats. The effect of ground nut oil in atheroge-
nic diets (i.e. cholesterol-containing) resembles that of a more saturated fat
but closer examination of the lesions shows that those due to saturated
fat are primarily lipid-rich 'foam' cells with little collagen deposition. The
typical lesions caused by ground nut oil are due to prominent intimal cell
proliferation associated with a particularly high collagen content, and
present as raised and sometimes very thick fibrotic areas (Kritchevsky et
al., 1971; Ehrhart and Holderbaum, 1980). Rhesus monkeys developed
progressive atherosclerosis with many of the morphological features of
human atherosclerosis (Vesselinovitch et al., 1974).
It has been postulated that arachidonic and behenic acids may contribute
towards the atherogenicity of groundnut oil. However, the removal of
these long-chain saturated fatty acids (which comprise 5-6% of groundnut
oil) in a specially formulated test oil of similar iodine value (105) to
groundnut oil did not prevent the formation of milder atherosclerotic
lesions (Kritchevsky et al., 1971). Therefore while atherogenicity may be
correlated with iodine value, some aspect of triglyceride structure must
also be important. Kritchevsky et al. (1981) showed atherogenic differ-
ences between groundnut oil from different geographic locations which
would suggest structural or compositional changes. Pancreatic lipase
specifically hydrolyses fats esterified to the 1 and 3 position of triglycer-
ides, which means lipid absorbed by the intestinal wall consists of
2-monoglyceride and free fatty acids. It has been shown that the triglycer-
ide structure of groundnut oil and the much less atherogenic randomized
groundnut oil differ in that the former contains more triglycerides with
linoleic acid (18:2) at the 2 position. While this alone would not explain its
atherogenicity over corn oil or the specially formulated test oil, there is the
possibility that more monounsaturated fatty acids in the 1,3 positions may
play an important role. Bezard and Sawadogo (1983) reported poor ab-
sorption into tissue triacylglycerols of long-chain saturated fatty acids in
comparison with monounsaturated oleic acid.
Randomized groundnut oil has a fatty acid composition identical to
groundnut oil but it has a different triglyceride structure and considerably
reduced atherogenicity. It has been shown to enhance cholesterogenesis in
vivo, which suggests it presents as being more unsaturated, despite having
the same iodine value as groundnut oil (Kritchevsky et ai., 1982).
Therefore the resulting micelle formed from the 2-monoglyceride, free
fatty acids and bile salts may differ in structure and composition, which
may in turn affect the structure of circulating lipoproteins (Kritchevsky et
ai., 1973; Kritchevsky et ai., 1984).
The ingestion of groundnut oil by chicks is accompanied by only
mild hypercholesterolaemia, even when fed with large amounts of chol-
esterol (Vesselinovitch et ai., 1974; Kritchevsky et ai., 1971). This is
in contrast to chicks fed cholesterol with hydrogenated groundnut oil. They
showed a massive increase in plasma cholesterol (Banerjee et ai., 1965).
An increase in the proportion of 13-lipoprotein as LDL (low density
lipoprotein) has also been noted (Kritchevsky et ai., 1976; Vesselinovitch et
ai., 1974).
It appears that the chemical nature of the lipid deposited in the
aortic wall may at least partly determine the quantity and quality of
the atherosclerotic lesions. While the influence of dietary fat on pre-
established lesions appears to be principally a function of the level of
unsaturation (Kritchevsky et ai., 1978), animals fed a cholesterol-free
diet with the addition of groundnut oil still showed the characteristic
lesions, although they contained relatively little lipid (Kritchevsky
et ai., 1971; Vesselinovitch et ai., 1974; Kritchevsky et ai., 1976).
The fact that the characteristic lesions were produced, even in the
absence of cholesterol, also strongly suggests a role for groundnut oil
triglycerides.
Studies of rapeseed (mustard) oil, which has a monounsaturated
fatty acid content similar to groundnut oil, have shown it to have a
similar or increased atherogenic effect, especially when the oils are
fed at high concentrations in the diet for an extended period of time
(Flanzy, 1979; Cluzan et ai., 1979; Sen and Gupta, 1979; Ray et ai.,
1980).
Protein 183
6.3.7 Absorption of fat
Levine and Silvis (1980) noted that the total fat in ground nuts was not
always efficiently absorbed in the gastrointestinal tract. Visual analysis of
stool samples taken from people fed whole groundnuts (76 g/day) showed
that portions of the nuts remained intact and were therefore not available
for lipid digestion (or any other nutrient). In these diets, 17.8% of the 80 g
of fat fed per day was found in the faeces. Simple grinding of whole
groundnuts (peanut butter) resulted in far less malabsorption of the fat in
the groundnuts (7.0% remained undigested). As expected, when the fat
was consumed as groundnut oil most was absorbed during its passage
through the tract (only 4.5% remained in the faeces).
6.4 PROTEIN
As more groundnuts are grown to meet the worldwide demand for oil,
greater amounts of groundnut protein are becoming available. This protein
is showing increasing potential as a food source, especially in the develop-
ing world where the lack of adequate protein can be a very serious dietary
problem. Research is leading to increased utilization of the protein, often
as part of processed foods.
Protein makes up 12.0-36.4% of the groundnut kernel (Hoffpauir, 1953;
Sekhon et al., 1970; Derise et al., 1974; Lusas, 1979; Woodroof, 1983). The
percentage increases during ripening, with the seeds from older plants
showing a higher capacity to accumulate protein than seeds from young
plants. The maximum increase occurs at 9 weeks; it then becomes approxi-
mately stable (Abdel Rahman, 1982b; Basha, 1991).
The storage proteins arachin and conarachin account for over 63% and
33% of the total protein content of groundnuts. The ratio of these globu-
lins ranges from 2: 1 to 4: 1, depending on the fractionation technique used.
Conarachin, found in the cytoplasm, has a basic nitrogen content of
6.5%, which is very high for seed globulins. It has a chemical score of
68-82% because of low levels of threonine (Thomson, 1952; Hoffpauir,
1953; Woodroof, 1983). Conarachin is a heterogeneous protein complex
with a high molecular weight. More recently it has been divided into three
subgroups, using DEAE cellulose chromatography. A major subfraction
a-conarachin has also been identified, and it is thought that this may
consist of two compounds whose molecular weights range from 140000 to
295000, depending on the ionic strength and pH of the buffer in which they
are dissolved.
Arachin, the second major storage protein, has a molecular weight of
330000 and is localized in the protein bodies ( aleurone grains) of the seeds.
It is much less soluble than conarachin and has a much lower chemical
score, due to limiting amounts of cystine and methionine. Polymorphism of
arachin has been reported (Tombs, 1965).
The lysine and methionine contents of conarachin I and II are signifi-
cantly higher than in arachin. A significant improvement in the overall
nutritive value of groundnut protein would be obtained if the proportion of
conarachin I and II, which is approximately 33% of the total protein
content, was increased at the expense of the arachin content. This is
especially important as it is quite clear that methionine and lysine are the
first and second limiting amino acids in groundnut protein.
A methionine-rich protein (MRP), consisting of six polypeptides with
molecular weights between 15.5 and 20 kD, has also been identified in
groundnut seed. Studies have shown considerable intervarietal variation in
the MRP composition, which shows the potential for improving the nutri-
tional value of groundnuts either by conventional plant breeding or by
genetic engineering (Basha, 1991). High performance liquid chromatogra-
phy has been developed as a rapid alternative to gel filtration in detecting
qualitative and quantitative differences in groundnut seed protein compo-
sition (Basha 1988).
6.4.1 Allergic reactions to groundnut protein

In some countries, ground nuts account for a significant proportion of the
dietary protein for both adults and children. Unfortunately groundnut
protein can cause immediate hypersensitivity reactions such as angioedema
and asthma in some individuals. Adverse reactions can range from abdomi-
nal discomfort to anaphylatic shock. Among foods that produce such
allergic reactions groundnut is considered second only to milk and eggs in
the total numbers of people affected.
Most individuals who are allergic to groundnuts avoid these problems by
eliminating groundnuts and groundnut products from their diets. In the
past this has not been a problem as groundnuts are usually easily recog-
nized in food. More recently identification has become a little more
difficult as groundnut flour is now added to a wider range of formulated
and processed foods.
Nordlee et al. (1991), using an in vitro radioallergosorbent test (RAST),
were able to show that groundnut oil, groundnut hull flour and a product
made from acid hydrolysis of groundnuts were not allergenic. Peanut
butter and products that had been processed to remove oil or heat treated
(roasted) were still allergenic. The remarkable heat stability of groundnut
allergens has been investigated by Neucere et al. (1969). Roasting of
groundnuts (145°C for 1 hour) produces little overall change in allergeni-
city as measured by RAST.
Barnett et al. (1983) were able to show that allergenicity is spread
through both crude arachin and conarachin fractions of groundnut protein
and that this activity was spread through the various molecular groupings
they were able to separate. Sachs et al. (1981) were able to isolate and
partially characterize a major groundnut allergen (Peanut-I) but they were
Protein 185
unable to suggest which major fraction of the storage protein it constituted.
They were able to show that Peanut-I was an acidic glycoprotein with non-
identical subunits. Peanut I appeared to be the most potent allergen they
isolated from groundnuts but not all of the allergenic activity in groundnut
extracts could be accounted for by this fraction.
The reason why groundnut protein appears to be so allergenic is due to
its remarkable thermal stability (Barnett et al., 1983) and its low protein
digestibility which means that it will remain in contact with the mucosa of
the small intestine for a longer time than a more easily digestible protein. It
is strongly recommended that individuals hypersensitive to groundnut
should avoid all products which may contain significant amounts of ground-
nut protein.
6.4.2 Amino acid composition

The amino acid composition of groundnuts is given in Table 6.3 with the
F AO reference protein for comparison While most of the amino acids are
found as part of protein and peptides, non-protein or free amino acids
(FAA) also occur. These are thought to react with glucose and fructose,
produced by the hydrolysis of sucrose during the browning process, to
produce the typical roasted groundnut flavour, colour and aroma (Wood-
roof, 1983; Young et al., 1974a). Specifically these are glutamic acid,
glutamine, asparagine, phenylalanine, aspartic acid and histidine. The
ratio of these amino acids alters with seed size, i.e. the different response
to roasting depends on the maturity of the seed (Young et al., 1974a).
Threonine and serine levels also increase with maturity (Young et al.,
1974b). Marshall et al. (1989) have developed a rapid method for the
determination of free amino acids in whole groundnuts.
High levels of free arginine, tyrosine, lysine and threonine have been
shown to affect the flavour adversely. Arginine and proline are found in
higher concentrations in immature kernels. Arginine levels decrease as the
kernels mature (Woodroof, 1983) but proline levels are more variable and
may be indicative of disease or other adverse conditions (Young et at.,
1974a).
The presence of two non-protein amino acids (,),-methyleneglutamine
and ,),-methyleneglutamic acid) in mature kernels has been confirmed
(Young et al., 1974b). Lee et at. (1965) have also reported an imino acid
derivative, N-methylhydroxyproline, in groundnut flour. At least six acidic
peptides have been reported in addition to the glutamic and aspartic acids
(Sundar et al., 1976). The role of these peptides in the seed has not been
established.
Table 6.3 shows that groundnuts are a reasonable source of essential
amino acids. Hoffpauir (1953) reported only lysine and methionine to be
deficient. Other authors suggest that isoleucine is also deficient (Evans and
Bandemer, 1967; Abdel Rahman, 1982a). Khalil and Chughtai (1983)
TABLE 6.3 Amino acid composition of groundnut
protein (g/lOO g kernels)
Groundnuts FAO a
Essential
Isoleucine 1.86-4.3 4.2
Leucine 6.12-7.0 4.8
Lysine 3,(1-4.27 4.2
Methionine 0.79-1.6 2.2
Methionine + Cystine 0.80-3.3 4.2
Phenylalanine 4.60-5.4 2.8
Threonine 2.43-2.9 2.6
Tryptophan 0.60-2.0 1.4
Valine 2.55-4.5 4.2
Non-essential
Alanine 3.4-4.2
Arginine 10.6-11.84
Aspartic acid 11.56-14.1
Cystine 1.3-2.42
Glutamic acid 19.2-22.46
Glycine 5.6-6.77
Histidine 2.1-2.51
Proline 4.3-6.36
Serine 4.9-5.30
Tyrosine 3.6-4.40
"FAO pattern of amino acid requirements (Evans and

Bandemer. 1967)
Based on data from: Hoffpauir. 1953; Evans and Bandemer,
1967; Chopra and Bhatia. 1970; Lusas. 1979; Alid et al .. 1981;
Sharma et al" 1981; Abdel Rahman. 1982a.
found only leucine, phenylalanine + tyrosine to exceed the reference

protein, in contrast to Sharma et at. (1981) who reported only phenylala-
nine. Like many other legumes, groundnut protein is also limited by the
availability of some essential amino acids. McOsker (1962) reported lysine,
methionine and threonine to be equally limiting while others report lysine
followed by threonine and methionine + cystine to be first limiting in
groundnut protein (Sharma et at., 1981; Khalil and Chughtai, 1983).
Methionine alone (Cheema and Ranhotra, 1967; Evans and Bandemer,
1967) or in combination with cystine (Evans and Bandemer, 1967) has also
been reported as the first limiting amino acid.
Most amino acids in proteins can be satisfactorily determined by the use
of an automatic amino acid analyser after acid hydrolysis of the protein.
Unfortunately tryptophan is destroyed during acid hydrolysis. A collabora-
tive study conducted by Westgarth and Williams (1974) showed that the
Protein 187
method devised by Miller (1967) using alkaline hydrolysis gave consistent
results for tryptophan on a standard groundnut sample, mean 0.98 g/
16g N. This value increased to a mean of 1.00g/16g N (range 0.91-1.07)
when the recovery of added tryptophan was considered. Amaya et al.
(1977) carried out an extensive survey of the tryptophan content of ground-
nuts grown commercially in the USA and reported that the tryptophan
content ranged from 1.05 to 1.41 g/16 g N. They acknowledged that the
tryptophan content of groundnuts was not nutritionally significant in the
light of the low levels of lysine and methionine. They noted an interesting
correlation between the tryptophan and total protein content among the
cultivated genotypes investigated. A wider variation in this ratio was found
when some wild genotypes were considered.
There appear to be considerable varietal differences in the total amino
acid content reported by various authors. These could, in part, be
accounted for by differing seed sizes and maturity as well as varying
methods of analysis. Other factors include location and environmental
conditions during growth, curing and storage. An example of the wide
variation of amino acid contents of a number of different groundnut
varieties in a single study is given by Heinis et al. (1975). Variation in
amino acids levels between different cuItivars of groundnuts should pro-
vide the opportunity for further genetic improvement. Glutamic acid,
aspartic acid and arginine account for approximately 45% of the total
amino acids; therefore a significant reduction of these three should pro-
duce significant increases in other amino acids (Young et al., 1973).
As lysine and methionine are the two main limiting essential amino
acids, various authors have determined the variables that may affect their
levels. Chopra and Sidhu (1965) attributed genetic similarity to the small
variation in total nitrogen, lysine and methionine levels found in their
study. In a further study Chopra and Sidhu (1967) showed a significant
difference (P < 0.01) in methionine levels between different spreading and
erect cultivars. Lysine levels were less significantly variable (P < 0.05).
Chopra and Bhatia (1970) showed no significant difference in the levels of
either of these amino acids due to habit or location of growth, but they
noted some intervarietal differences. These were more marked for lysine
(2.67-4.27%) than methionine (0.79-0.96%).
In contrast, a study of three cultivars (virginia, red and white skinned
spanish) by Conkerton and Ory (1976) showed no significant difference in
lysine or methionine levels (3.1-3.3% and 0.9-1.1% respectively). This is
supported by Young (1980) who studied the three types, spanish, virginia
and runner. He reported a significant location effect for alI the amino acids
but variety had no effect on methionine, lysine or arginine levels. Looking
at cultivars of the spanish type, Young et al. (1974a) showed intervarietal,
location and environmental (irrigation) differences to be considerable for a
number of amino acids while lysine and methionine levels were unaffected.
Similarly, habit and location of growth showed no significant influence on
methionine levels in a study by Cheema and Ranhotra (1967). These
variables seem to play a more significant role in the levels of the amino
acids related to organoleptic qualities rather than nutritive quality (Young
et al., 1974a). However, Young et al. (1973) investigated the amino acid
levels of defatted groundnut meal produced from 16 different varieties of
groundnuts with widely differing protein content. They found nearly two-
fold variations in the limiting essential amino acids (lysine, isoleucine,
methionine, threonine and valine) - large enough to be significant for
breeding purposes. From this it could be interpreted that varietal differ-
ences are linked to total nitrogen levels.
Amaya et al. (1977) investigated the correlation between tryptophan
levels and total protein as a means of indicating the abundance of other
essential amino acids. They found the former tends to remain constant
while the total protein varies substantially with different environmental
conditions. Sharma et al. (1981) demonstrated a marked change in protein
quality in mutants produced by gamma-irradiation. They showed lower
sulphur amino acids levels and a higher tryptophan content compared to
the parent.
6.4.3 Processing of groundnuts and their products

Often processing only involves roasting prior to consumption as a snack
food or grinding into peanut butter. Some authors report that the overall
protein content of groundnuts appears to be little affected by roasting
(Sekhon et al., 1970; Woodroof, 1983) while Derise et al. (1974) and Khalil
and Chughtai (1983) reported a slight increase, probably because of an
overall decrease in weight due to loss of moisture and non-nitrogenous
volatiles. In many groundnut producing countries, the protein-rich meal
that remains after oil extraction is used as animal feed or fertilizer. This is
now being recognized as a valuable supplement for human diets. Further
processing provides flour, protein concentrates and isolates.
Groundnut flour can range from full-fat to defatted, with the protein
level increasing as the fat level falls. The protein content of defatted
groundnuts is reported to range from 43.8% to 68.6% (Howe et al., 1965;
Cheema and Ranhotra, 1967; Fetuga et al., 1973; Ayres et al., 1974;
Bookwalter et al., 1978, 1979; Ory and Conkerton, 1983; Khalil et al.,
1983b; Alid et al., 1981; Ghuman et al., 1990). Screw pressing or pre-
pressing followed by solvent extraction are the two main commercial
methods for producing the defatted product. The latter then requires the
solvent to be stripped from the meal by steam heating, which also removes
the raw groundnut flavour (Natarajan, 1980). As with other processes, the
heating stage is critical; severe heat can affect the availability and utiliz-
ation of the essential amino acids. Ayres et al. (1974) found the nitrogen
solubility index fell from 92% in defatted raw groundnuts to 59% for
pre-pressed solvent extracted flour.
Protein 189
Further processing of the meal to produce protein concentrates and
isolates has the advantage of removing the insoluble and partly indigestible
carbohydrates which constitute approximately one third of the raw
material. Groundnut protein concentrate (PPC) , containing 65-70%
protein, can be produced by wet-milling, which gives simultaneous recov-
ery of oil and protein (Natarajan, 1980). Khan et at. (1975) reported levels
of 54.3-56.1%. Alternatively concentrates can be prepared by leaching
meals with dilute acids and aqueous alcohols to give a product with higher
protein content, considerably lower trypsin inhibitor content and negligible
aflatoxin levels. The nutritional quality of these concentrates remain un-
affected but the protein solubility is decreased, owing to denaturation that
occurs during extraction (Nagaraj and Subramanian, 1974).
Groundnut protein isolates are the most refined form of groundnut
proteins, going one step further than the concentrate process and removing
water-insoluble polysaccharides, water-soluble sugars and other minor
constituents (Anantharaman et at., 1969; Bhatia et at., 1960; Natarajan,
1980). Depending on the method of isolation, they contain 90% (wet-
milling) to 95% (solvent extracted meal or screw press) protein.
The overall effect of processing groundnuts to produce flour, concen-
trate and isolates is to increase markedly the crude protein and conse-
quently the essential amino acid content by the removal of the
carbohydrate fractions (Table 6.4).
Fermentation of groundnut kernels or defatted groundnut flour (DPF)
supposedly enhances the digestibility by breaking down the complex
TABLE 6.4 Essential amino acids levels in groundnut products (g/lOO g)
Flour Concentrate Isolate
Essential
Isoleucine 1.64-3.4 3.5-4.3 3.6
Leucine 3.64-6.79 6.7-7.09 6.6
Lysine 2.70-4.55 2.9-3.9 3.0
Methionine 0.51-1.21 0.9-1.1 1.0
Methionine + Cystine 1.8 2.05
Phenylalanine 2.69-5.54 5.5-5.6 5.6
Phenylalanine + Tyrosine 4.9 9.94
Threonine 1.46-3.4 2.5-3.04 2.5
Tryptophan 0.55-1.0 0.9-1.24 1.0
Valine 2.18-4.1 4.27-4.7 4.4
Non-essential
Cystine 1.97-3.31 1.14-1.4 1.4
Tyrosine 3.36-3.96 4.1-4.34 4.3
Based on data from: Young et al., 1973; Fetuga et al., 1973; Ayres et al., 1974; Khan et al.,
1975; Conkerton and Ory, 1976; Bookwalter et al., 1978; Lusas, 1979; Alid et al., 1981.
protein structures to pep tides of varying lengths and free amino-acids,
which in turn increases the nutritive value (Coelho and Bhat, 1959; Rao
and Rao, 1972; Quinn et at., 1975; Bhavanishankar et at., 1987).
Alternatively, proteolytic enzymes can be used to modify and extract
protein from groundnut flour and press cake (Natarajan, 1980). The prod-
uct is known as hydrolysed vegetable protein. Texturized proteins have
also been developed from DPF (Alid et at., 1981).
6.4.4 Availability of amino acids

The key to amino acid availability in groundnuts and their products lies in
the processing they undergo before consumption. Raw groundnuts contain
a number of anti nutritive factors including trypsin inhibitors, haemaggluti-
nins, goitrogens, saponins and phytic acid. While the level of trypsin
inhibitor found in groundnuts is only approximately 20% of that found in
raw beans, it is still enough to cause pancreatic hypertrophy in rats
receiving 15% protein intake from groundnuts (Lusas, 1979). Groundnuts
are unusual in that, while most of the protein is concentrated in the outer
cotyledon layer, the antitrypsin activity is concentrated in the inner part of
the cotyledon (Zimmermann et at., 1967). Mild heat treatment, an in-
herent part of most of the processes, is sufficient to destroy the trypsin
inhibitor and haemagglutinin in the seeds.
Mild dry heating increases available lysine and therefore protein
efficiency ratio (PER) values but continued high-temperature roasting
decreases the protein solubility and protein quality. Roasting has been
reported to decrease the measurable free amino acid content by 77.8%
(dry), 64.6% (microwave) and 52% (oil) (Lusas, 1979). Aspartic acid,
serine, glutamic acid, proline and phenylalanine levels increase while
threonine, tyrosine, lysine, arginine, methionine, cystine and tryptophan
levels all fall, with lysine, methionine and threonine suffering maximum
loss (in that order) (Khalil and Chughtai, 1983). McOsker (1962) also
found these three to form the limiting amino acid sequence in roasted
groundnuts but in a slightly different order. He found roasting at 170°C for
36 minutes resulted in the destruction of 15% lysine, 11% threonine and
10% methionine. Similarly Woodroof (1983) reported a decrease in lysine
levels from 3.4% to 1.9% for meal prepared by cooking at 250 OF for 2
hours.
Anantharaman and Carpenter (1971) looked closely at the effects of
heat and found that higher temperature, longer heating time and higher
moisture content all resulted in reduced values for each amino acid
investigated. Autoclaving at 107°C for 30 minutes (similar to heating in
commercial groundnut meal production) has little effect on cystine and
lysine levels. More drastic heating (121°C for 4 hours) reduces FDBP-
available lysine by nearly 40% and total cystine by nearly 30% while
appearing to have little effect on available methionine and threonine
levels. The reduction in E-NH2 lysine groups is attributed to binding to
Protein 191
hydrolysed sucrose. Linkages are formed which are not hydrolysed by
digestive enzymes.
Quinn et al. (1975) found that fungal fermentation of DPF gave a slight
decrease in arginine and proline levels while glycine and alanine seemed to
be slightly increased. They also noted slight increases in lysine and meth-
ionine levels. Bhavanishankar et al. (1987) noted certain fungal strains
have higher proteolytic activity, resulting in increased free lysine and
methionine levels. In contrast Sugimura et al. (1965) found considerably
increased levels of free amino acids, especially proline and glutamic acid,
while arginine levels decreased. The samples were autoclaved in each of
these studies prior to fermentation.
The available lysine content of groundnuts is high (2.69 g/16 g N) com-
pared with the levels found in cereals (Johri et al., 1988) but on long-term
storage (up to 24 months) the levels can fall considerably, especially at high
temperatures (40°C). Similar falls in available lysine occurred even when
they were stored in a vacuum.
6.4.5 Protein quality

The nutritional quality of protein depends on the amount and kind of
essential amino acids which become available during digestion. If anyone
of the essential amino acids is deficient, it becomes the limiting factor in the
utilization of that protein. Completeness of digestion and absorption of the
essential amino acids is also important (Watts et al., 1959). The nutritional
value has been assessed for a number of different groundnut products, e.g.
meal, flour or defatted oil cake. The use of heat in the processing of these
products can reduce the solubility of the protein as well as destroying some
of the sulphur-containing amino acids (Ladell and Phillips, 1959). This
variable alone may account for some of the variability of results that have
been reported.
In vivo biological evaluation to determine the biologi,cal value (BV)
(Watts et al., 1959), protein efficiency ratios (PER) (Howe et al., 1965;
Nagaraj and Subramanian, 1974; Ghuman et al., 1990) and net protein
utilization (NPU) (Khalil and Chughtai, 1983) are commonly employed to
evaluate protein quality. A summary of some typical values for groundnut
and groundnut products are shown in Table 6.5. Alternatively, microbio-
logical techniques can be used for the rapid testing of protein quality.
These measure the growth of the organism in question (usually
Tetrahymena pyriformis W) to give the relative nutritive value (Ghuman et
al., 1990).
Because groundnuts are deficient in some of the essential amino acids
(notably lysine and methionine), their relative nutritive value is decreased
in comparison with a reference protein. They have a chemical score of
between 58 and 70 (Khalil and Chughtai, 1983). Cheema and Ranhotra
(1967) used the chemical score method to assess the biological value of
TABLE 6.5 Comparisons of protein quality for raw
and processed groundnuts
Raw Roasted!
boiled
defatted
True digestibility 89-93

Biological value 32-75
Nutritive value 47.2-54.2
Protein efficiency ratio 1.5-1.8 1.45-1.76
Net protein utilization 34-44
Available lysine 2.83%
Based on data from: Hoffpauir, 1953; Clegg, 1960; Howe et

ai., 1965; Evans and Bandemer, 1967; Nagaraj and
Subramanian, 1974; Lusas, 1979; Woodroof, 1983; Alid et ai.,
1981; Khalil and Chughtai, 1983; Ghuman et ai., 1990.
defatted groundnuts which ranged from 51.3 to 58.0. Biological value is

more often determined by incorporating the protein (usually at 10%) in a
test animal's diet. The reported BV for raw groundnuts is 50-75% of the
reference proteins and the PER is 1.5-1.8 (Lusas, 1979). Studies have
indicated that the digestibility and utilization of ground nut protein from
kernels, shown by nitrogen balance, is poor (Watts et al., 1959; Hoffpauir,
1953; Edwards et al., 1966; Woodroof, 1983). This is probably due in part
to the presence of a trypsin inhibitor and fibre. Khalil and Chughtai (1983)
compared NPU from several strains after roasting and found the differ-
ences in nutritional quality (34 to 44) were minimized by the higher
amounts of lysine in the strains containing the lower amounts of total
protein.
Coelho and Bhat (1959) compared the digestibility, BV and PER of raw
vs roasted/boiled kernels after fermentation. While the levels of digestibi-
lity were not significantly different, the BV increased from 56% to 75.5%
and 76.4% respectively after boiling and fermentation. Roasting appeared
to be the least effective treatment, resulting in a BV of 69.6%. In contrast
the PER values for all groups were comparable (1.14-1.38) which suggests
that, while the fermentation of groundnuts may help in an improved
utilization of their proteins, it contributes nothing towards digestibility or
growth. Bhavanishankar et al. (1987) found a significant difference in the
PER of two different strains of fungi (1.36 and 1.75) and this suggests that
one important factor in fermentation is the fungal strain used.
Defatted groundnut flour (DPF) shows little nutritive difference com-
pared with raw groundnuts, with PERs ranging from 1.45 to 1.76 (Nagaraj
and Subramanian, 1974; Howe et al., 1965; Alid et al., 1981; Ghuman et
al., 1990). Mild heating of the flour has little effect, but autoclaving for 4
Protein 193
hours at 121°C reduced its protein value from 60% to 53.9% (Ananthara-
man and Carpenter, 1971). Ghuman et al. (1990) report that the chemical
score for S-amino acids and lysine ranges from 37% to 50% and from 47%
to 55% respectively, with a significant and positive correlation between
PER and methionine levels. They also found highly significant correlations
between conarachin values and the values of the two limiting amino acids,
methionine and lysine. This suggests that cultivars with a high conarachin
content are better for breeding purposes. Evans and Bandemer (1967)
report a protein nutritive value (weight gain x 100/weight gain of rats fed
casein) ranging from 32% to 49%. A similar assay utilizing Tetrahymena
pyriformis W found a range from 45.6% to 54.2% (Ghuman et al., 1990).
This result was significantly linked to the chemical score and PER and can
be used as a quick method to measure protein quality.
The limited availability of lysine and methionine are linked to decreased
BV and PER compared with the WHO/FAO reference protein. Authors
differ in their views as to the effects of supplementing the groundnut
protein with the deficient essential amino acids. These differences may in
part be accounted for by the varying amounts added. Howe et al. (1965)
reported lysine, methionine and threonine to be equally limiting in a diet
containing 10% DPF. They used 0.4% threonine, 0.2% lysine and 0.2%
methionine to give a PER (2.22) close to that of casein (2.5). Alid et al.
(1981) reduced the threonine to 0.3% (which still exceeded the WHO/
FAO reference pattern for essential amino acids), with a resulting PER of
2.18. McOsker (1962) showed that supplementation of roasted groundnuts
with at least 0.31% L-Iysine, 0.19% DL-threonine and 0.21% DL-
methionine showed feed efficiency equal to or better than that of a 15%
casein diet.
Evans and Bandemer (1967) showed no effect with supplementation of
10% groundnut protein with 0.3% methionine, 0.1 % isoleucine and 0.1 %
lysine. They postulated the need for threonine as well. In another experi-
ment, they measured bacterial growth, this time increasing the methionine
to 3.3%, isoleucine to 0.7% and the lysine to 0.5%. The protein nutritive
value increased from 26% to 81 %. Adding 0.2% valine and 0.2% threo-
nine caused this to decrease to 45%. Joseph et al. (1974) reported that the
addition of 1.5% L-Iysine and 1% DL-methionine to a diet containing
groundnut protein at a dietary level of 10% increased the PER from 1.65 to
2.07 after 4 weeks. Addition of 1% threonine further increased this to 2.6.
The supplementation of an extrusion texturized. protein with 0.3% DL-
threonine, 0.2% L-Iysine and 0.2% DL-methionine gave a similar PER
(2.18) to the similarly supplemented starting product (DPF) , which
suggests that there is no important change in the protein quality during this
process (Alid et al., 1981).
Quinn et al. (1975) found that feeding fermented groundnut flour
(at 15%), supplemented with 0.4% methionine, 0.25% lysine and 0.13%
tryptophan, made no difference to the PER (2.37-2.44) in comparison with
supplemented, non-fermented flour (2.5-2.53). It would be interesting to
know the PER values for unsupplemented flour. Anantharaman and
Carpenter (1971) found supplementation of both unheated and mildly
heated DPF improved the PER, the order of limitation being methionine
> lysine > threonine. Severe heating of the flour changed this order to
lysine> cystine> methionine> threonine.
It was thought by some that raising the dietary level of groundnut
protein might increase its biological usefulness. Lusas (1979) reported rats
fed 16.7-20% groundnut protein grew as well as those fed 12-24% casein;
however, Murphy and Dunn (1950) showed a diet containing 19% ground-
nut protein needed to be supplemented with 1.6~!" lysine plus 0.58%
methionine to be adequate for lactation and reproduction. Lysine supple-
mentation alone was insufficient.
It is important to note that treatment of the groundnut meal or flour to
reduce aflatoxin contamination can markedly reduce the protein quality,
reflected in lower PER values (Ihekoronye, 1987). However, it is still
suitable for supplementing high carbohydrate, cereal-based foods.
6.4.6 Supplementation of diet

Protein quality is often assessed on the 'completeness' of the single protein,
rather than its supplementary value in a mixture from a variety of plant and
animal sources. More recently the use of groundnut meal is becoming more
recognized, not only as a dietary supplement for children on protein-poor,
cereal-based diets in economically underdeveloped countries, but also as
an effective treatment for children with protein malnutrition, i.e. kwash-
iorkor (Rao et al., 1965a; Joseph et al., 1960). The nutritional value can be
improved by combining it with an animal protein such as skim-milk powder
or plant proteins which complement it, or by supplementing it with the
deficient essential amino acids (Waldo and Goddard, 1950; Sanchez et al.,
1972; Joseph et al., 1974). Rao et al. (1965) showed that, while the protein
content remained the same (49.2), supplementing 50% groundnut flour
with 50% full-fat soya flour and 1% each of L-Iysine and DL-methionine
increased the PER to 2.8 and NPU to 72.3.
Protein-rich groundnut flour can be used to supplement corn or wheat
flour for the production of a more nutritious bread, a staple food through-
out the world (Khalil et al., 1983a; Khalil et al., 1983b). Supplementing
wheat flour with 30% groundnut flour gave a 100% increase in protein
content of the bread. However, the best result in terms of PER, NPU and
organoleptic evaluation was achieved when the bread was supplemented
with 20% groundnut flour (Khalil et al., 1983a). Ory and Conkerton (1983)
found that the use of more than 12.5% protein flour required dough
conditioners to prevent a decrease in loaf volume during baking.
While supplementation of bread with protein flour increases the overall
Protein 195
protein content and generally results in a more positive nitrogen balance
(due to a better balance of the essential amino acids), increased fibre
content can lead to decreased digestibility of the product (Khalil et ai.,
1983b). Therefore while the PER might increase in the supplemented
product, it may still fall a long way short of the reference protein. Khalil et
ai. (1983a) showed the PER of bread increased from 1.21 to a maximum
1.61 (supplemented with 20% groundnut flour) while the NPU increased
from 62% to 80% at this level. Substitution of one half of the groundnut
flour with either soy or chick-pea flour has been shown to improve the
nutritive quality (Bookwalter et ai., 1978; Khalil et ai .. 1983b). Bread made
with groundnut protein concentrate showed superior flavour, taste and
crumb colour but concentrations above 10% reduced loaf volume consider-
ably (Khan et ai., 1975). Available lysine does increase with supplemen-
tation, but it is also lost during cooking (Bhat and Vivian, 1980).
Bookwalter et ai. (1978), using 15% defatted groundnut flour, increased
the PER from 1.4 to 2.0 with the addition of 0.18% L-lysine. No further
improvement was noted with increased lysine levels (Bookwalter et ai.,
1979). Alternatively, the addition of 20% skim-milk powder substantially
increases the PER (Waldo and Goddard, 1950; Joseph et ai., 1974).
However, if these dietary supplements are further processed by cooking
(e.g. as biscuits made with groundnut flour, wheat flour and skim-milk
powder) the available lysine content is reduced, which in turn lowers the
value of the protein (Clegg, 1960).
Children with kwashiorkor fed experimental blends of two thirds
groundnut protein isolate combined with one third dry skim-milk (to give
an average protein content of 69%) or 50% isolate with 50% casein plus
2% lysine and 1% methionine (to give an average protein of 83.6%) gave
a similar clinical response to those fed skim-milk (Webb et ai., 1964).
The isolate contained approximately 90% protein. Shurpalekar and
Bretschneider (1969) found only 22% of isolate was required to make
biscuits with a protein content of 28.8%, suitable for the prevention and
treatment of malnutrition. These biscuits had good organoleptic and keep-
ing qualities in comparison with biscuits made with defatted groundnut
flour, which were reported to be hard (Ory and Conkerton, 1983).
Alternatively, the protein isolate can be combined with lentil flour, raw
sugar and hydrogenated groundnut oil, to be eaten as a sweet (Dumm et
ai., 1966).
Groundnut protein, once used only as animal feed after the extraction of
oil from kernels, is gaining credibility as a readily available source of
protein to meet world demand. While invariably deficient in the essential
amino acids lysine, methionine and threonine, the protein quality is of a
sufficient standard to increase protein efficiency ratios when added to a
staple cereal diet. Processing of the meal to remove carbohydrates and
fibre increases the digestibility and protein content of the final product.
Supplementation of the deficient amino acids, with either another plant
protein or the deficient amino acids, can provide a food capable of meeting
dietary requirements even in the malnourished.
The quality of groundnut protein would also be improved if the levels of
the three main limiting amino acids were increased in the seed protein. If
plant breeders could identify cultivars that consistently produce higher
levels of any (or all) of these three, regardless of location or environmental
effects, the resulting protein would have even more value.
6.4.7 The effect of storage on protein quality

Yannai and Zimmermann (1970) noted that storage of groundnut meal for
6-18 months at relative humidities of 40% and 60% resulted in a slight
drop in the nutritive value of the meal. The PER of the meal increased on
storage to 24 months and at some storage temperatures was greater (3.19)
than the PER value of the freshly prepared meal (2.84). It is interesting to
note that the trypsin inhibitor activity of the unheated groundnut meal
decreased significantly during this storage trial. The samples stored at the
highest temperatures showed greater reductions. It is quite possible that
the increases in nutritive value observed by Yannai and Zimmermann
(1970) were a direct result of trypsin inhibitor degradation. It should be
noted that the successful storage of this meal at adverse temperatures and
relative humidities for up to two years was a direct result of the efficient
removal of oil from the de hulled seeds at the commencement of the
experiment.
6.5 CARBOHYDRATE CONTENT
Carbohydrates in groundnut kernels consist of water-soluble carbo-

hydrates (monosaccharides, disaccharides) and oligosaccharides (including
starch, raffinose and stachyose). These together comprise the total avail-
able carbohydrate fraction. A summary of typical values is shown in Table
6.6. Two reviews of ground nut carbohydrates with good summaries of the
non-starchy polysaccharide fraction have been published by Tharanathan
et al. (1975, 1979).
Carbohydrate levels are high in newly germinated groundnut kernels but
the levels begin to fall from an average of 46% at 5 weeks as the maturing
seed begins to store oil (Abdel Rahman, 1982b). Mature kernels are
reported to contain 9.5-19.0% total available carbohydrate as both soluble
and insoluble carbohydrate (Rao etal., 1965a; Oke, 1967; Abdel Rahman,
1982b; Woodroof, 1983). Water-soluble carbohydrates comprise 3.0 to
8.9% of the available carbohydrate (Sekhon et al., 1970; Wallerstein et al.,
1989; Rodriguez et al., 1989). Seasonal variation plays a role in the levels of
both soluble and insoluble carbohydrates. Rodriguez et al. (1989) analysed
soluble carbohydrate levels from a crop on two successive years. The first
Carbohydrate content 197
TABLE 6.6 Carbohydrates in groundnut kernels and flour (%)
Kernels Flour
Soluble sugars
Monosaccharides 0.1-0.3 0.13
Sucrose 1.9-5.2 7.70
Total water-soluble carbohydrates 3.0-8.92
Oligosaccharides
Raffinose 0.14
Stachyose 0.71
Starch 6.7-10.0
Total available carbohydrates 9.5-18.6 22.1-32.85
Insoluble carbohydrates
Total insoluble carbohydrates 12.2-15.67
Crude fibre 1.2-4.3 2.7-5.76
Neutral-detergent fibre 8.3-11.2 8.4
Acid-detergent fibre 4.3-5.9
Lignin 1.1-3.3
Based on data from: Anantharaman et al .. 1959; Ladell and Phillips. 1959; Howe el al .• 1965;
Rao et al .. 1965a; Oke. 1967; Daghir et al .. 1969; Yannai and Zimmermann. 1970; Ayres et
al .. 1974; Bookwalter et al .. 1979; Alid et al .. 1981: Woodroof. 1983; Ory and Conkerton.
1983; Wallerstein et al .. 1989; Rodriguez et al .. 1989.
year ranged from 7.14% to 8.92%; the following year's crop ranged from
5.49% to 7.82%. Analysis of insoluble carbohydrates showed little differ-
ence between the two crops (12.79% to 15.67% compared with 12.18% to
15.43% ).
The amount of total available carbohydrate falls slightly if the ground-
nuts are skinned, whereas boiling can account for a loss of 4% (Woodroof,
1983). Water-soluble carbohydrates comprise 3.0 to 6.17% of the available
carbohydrate (Sekhon et ai., 1970; Wallerstein et ai., 1989). Roasting the
kernels causes an increase in available carbohydrate, probably due to the
loss of volatiles and water. It also results in the hydrolysis of sucrose, the
predominant water-soluble carbohydrate, to glucose and fructose. These
then react with free amino acids (Maillard reaction) to give the character-
istic flavour of roasted groundnuts (Young et ai., 1974b). This is supported
by Sekhon et ai. (1970) who reported an increase in reducing sugars, offset
by a decrease in non-reducing sugars and total water-soluble carbohydrate
during roasting. Differing soluble carbohydrate levels, due to varietall
environmental differences or level of maturity, may all contribute to
variation in roast groundnut kernel characteristics (Rodriguez et ai., 1989).
Carbohydrate levels in groundnut products depend on the method of
processing used (Table 6.6). Groundnut flour, made from defatted ground-
nuts, contains protein and the soluble and insoluble carbohydrates.
Groundnut protein concentrate also contains some insoluble carbo-
hydrates, whereas ground nut isolates should only contain protein (Natara-
jan, 1980).
The total carbohydrate level in groundnut flour is reported to range
between 22.11 and 32.85% (Anantharaman et al., 1959; Daghir et al.,
1969; Yannai and Zimmermann, 1970; Fetuga et al., 1973; Ayres et al.,
1974; Conkerton and Ory, 1976; Bhat and Vivian, 1980; Alid et al., 1981;
Carew et al., 1988; Woodroof, 1983).Sometimes reported as nitrogen-free
extractives (NFE), it is usually estimated by the difference in proximate
analysis, which can account for the wide range in reported values. For
example, Rao et al. (1965) found 18.5% carbohydrate in a 1:1 blend of
ground nut and soya flour, compared with 28.9% found by Shurpalekar and
Bretschneider (1969). The other major difference between the two was in
moisture levels - 7.2% for the former compared with 2.9% for the latter.
Ostrowski et al. (1971) reported groundnut concentrate with 24.1%
nitrogen-free extractives, a similar level to that found in groundnut flour,
whereas Khan et al. (1975) gave a figure of 18.5%.
The starch levels in groundnut flour inhibit the formation of extrusion-
texturized protein. They tend to gelatinize upon extrusion and disrupt the
protein matrix necessary for texture formation (Alid et al., 1981). This can
be controlled by the addition of surfactants.
Whole-wheat flour has a carbohydrate content which ranges from 72.1 %
to 81.2%. The addition of groundnut flour to wheat flour decreases the
total carbohydrate levels while increasing protein content (Bhat and
Vivian, 1980; Khalil et al., 1983b; Ory and Conkerton, 1983). In contrast,
products such as muffins made from 100% groundnut flour had a carbo-
hydrate level of 23.2% (Ory and Conkerton, 1983). Tharanathan et al.
(1976) analysed the soluble carbohydrate composition of defatted ground-
nut flour. Glucose was the most predominant (2.9%), followed by fructose
(2.2%) and sucrose (0.9%). Raffinose and stachyose levels were low
(0.94% and 0.20% respectively). Since they are both poorly digested and
responsible for flatulence in humans, low levels make the groundnut flour
more acceptable as a food supplement than some other plant proteins, such
as soybean flour which can contain higher levels of these indigestible
oligosaccharides.
The fermentation of groundnut meal initially causes a slight fall in
carbohydrate levels (after 18 hours), which then increases again. It is
thought that the increase may be due to degradation of the complex
carbohydrates, such as raffinose and stachyose (Bhavanishankar et al.,
1987). Quinn et al. (1975) used a 4-day fermentation period and found the
carbohydrate levels fell from 35.1-35.8% to 29.3-33.2%. The variation
was attributed to the different strains of fungi used for the fermentation,
some of which hydrolyse stachyose and/or raffinose better than others. The
preparation of a miso-like product, which involves two separate and dis-
tinct fermentations, resulted in an increase in reducing sugars from 4.34%
to 8.57% after 20 days fermentation (Rao and Rao, 1972).
Carbohydrate content 199
6.5.1 Fibre
Plant fibre is generally resistant to hydrolysis by human digestive enzymes
and remains after food has passed through the small intestine. As such it is
referred to as 'unavailable carbohydrate', even though it undergoes some
bacterial degradation in the colon. The fibre content of a food can lead to
decreased digestibility of the proteins, therefore reducing amino acid
availability. Groundnuts contain fibre which consists principally of the
carbohydrates cellulose, hemicellulose and pectin. This fraction also con-
tains smaller amounts of mucilages, gums and lignin. Wallerstein et al.
(1989) report that the lignin content of groundnuts ranges from 1.1% to
3.3% for three cultivars. It should be noted, however, that lignin is not a
polysaccharide - it is just found associated with the fibre fraction. The
terms crude fibre, acid-detergent fibre and neutral detergent fibre denote
different methods to estimate different fractions present in food. While the
estimation of crude fibre seems to be widely used, it is not a true indication
of total dietary fibre (Ory and Conkerton, 1983).
Crude fibre levels in raw groundnuts are reported to range from 1.2% to
5.0% (Rao et al., 1965a; Oke, 1967; Derise et al., 1974; Woodroof, 1983).
This figure increases slightly for acid-detergent fibre (4.3% to 5.9%), and
significantly for neutral-detergent fibre (8.3% to 11.2%) (Wallerstein et
al., 1989). Fibre content decreases slightly with boiling or removal of skins,
whereas roasting leads to a slight increase (Woodroof, 1983; Derise et al.,
1974).
The reported levels of crude fibre for groundnut flour are similar:
2.70-5.76% (Anantharaman et al., 1959; Ladell and Phillips, 1959; Howe
et al., 1965; Woodroof, 1969; Natarajan, 1980; Daghir et at., 1969; Yannai
and Zimmermann, 1970; Fetuga et al., 1973; Ayres et al., 1974;
Bookwalter et al., 1979 Alid et al., 1981). In contrast to this, Carew et al.
(1988) give a figure of 10.6% for crude fibre in groundnut meal - almost
double the amount - which may be attributable to the different methods
used to measure this fraction. Crude fibre in groundnut concentrate ranges
from 4.0% to 6.3% (Ostrowski et al., 1971; Khan et al., 1975).
The fibre content of a flour is increased if it is made from unskinned
groundnuts. Ory and Conkerton (1983) calculated the neutral-detergent
fibre (NDF) level of groundnuts and skins to be 8.4%, whereas the NDF of
the skins alone ranged from 35.1 % to 41.1 %. As such, the skins contribute
to the total fibre content of the flour. This could be exploited in developed
countries where levels of dietary fibre are often low. The addition of 12.5%
groundnut flour in breads increases the dietary fibre content from 0.3%
(wheat) to 0.4-0.6% (Ory and Conkerton, 1983). Kahn et al. (1975)
attained slightly higher levels with 15% DPF (0.7-0.8%) and 15% ground-
nut protein concentrate (0.9%). Muffins, made from 100% groundnut
flour, attained fibre levels of 2.2% COry and Conkerton, 1983).
Fermentation of groundnut meal for up to 22 hours showed no change in
the crude fibre content of any of the samples (Bhavanishankar et at., 1987).
Quinn et at. (1975) reported a slight increase in crude fibre content of the
fermented samples, perhaps due to the loss of volatiles during fermen-
tation (this process took 4 days at 28°C).
A compound not classified as fibre but often found associated with it in
foods is phytic acid. This is present as phytin (the mixed calcium, mag-
nesium and potassium salt of phytic acid) in groundnut flour at a concen-
tration of 3.2%. Phytic acid is able not only to bind calcium in the diet and
reduce availability but also to bind other useful elements such as zinc and
magnesium. Groundnut kernels contain 3.3% of the total phosphorus as
phytic acid phosphorus, so that this is also unavailable (Oke, 1967).
6.6 VITAMINS
Raw groundnuts are known to be an excellent source of certain vitamins,

especially E, K and the B group. The groundnut is one of the richest
sources of thiamin (Bj) in plants. Dougherty and Cobb (1970a,b) noted
that 90% of the total thiamine content of groundnuts was contained in the
testa, principally in the cell wall fraction. They also noted that approxi-
mately 11 % of this migrated into the cotyledons when they were dried.
Chaturvedi and Geervani (1986) noted that groundnuts were also an
excellent source of niacin which ranged from 16.7 to 21.7 mg/lOO g DM
basis for a number of Indian cultivars. They observed that boiling resulted
in a small loss of niacin, while roasting and frying resulted in a small
increase due to loss of moisture. In human experiments they observed that
91.7% of the niacin in boiled groundnuts was available, falling to 87.7%
for roasted groundnuts. Chaturvedi and Geervani (1986) suggest that the
niacin content of groundnuts makes a significant contribution to the diets
of people living in dryland areas. This is important as diets in these areas
contain limited amounts of the essential amino acid tryptophan; niacin,
however, in adequate amounts can spare tryptophan for protein synthesis.
Rao et at. (1965b) noted that the tocopherol content of Indian groundnut
oil was 0.93 mg/g oil and it was made up of 36% a and 64% 'Y tocopherol.
The total amounts of tocopherol found in groundnut oil were similar to
those found in other locally produced oils and to imported soya bean oil.
The colour of groundnut oil is in part due to the carotenoid content. It is
well known that there is a marked reduction in the colour of groundnut oil
as the seed matures. Pattee et at. (1969) showed that this reduction in
colour was due to a rapid increase in the oil content of the seed which
diluted the carotenoid content. Pattee and Purcell (1967) showed that a
sample of oil extracted from immature virginia groundnuts contained 60 I-1g
l3-carotene and 138 I-1g lutein/litre of oil. The total concentration of carote-
noids from mature groundnuts was less than 1l-1g/litre of oil.
Mineral composition 201
6.7 MINERAL COMPOSITION
The total ash content of groundnuts and products derived from groundnuts
is shown in Table 6.7. The rise in the total ash content of defatted
groundnut flour is expected as fat is removed from the kernels; in the same
way a reduction in the total ash content of protein concentrates or isolates
is to be expected as these are easily removed in the washing process. Abdel
Rahman (1982b) noted that the total ash content ranged from 1.95% to
2.10% in the growing seed in the developmental period from 5 to 12 weeks.
The mineral content of raw groundnuts and some of their products is
shown in Table 6.8. In general, groundnuts provide a wide range of
mineral elements to supplement human dietary requirements. They have
the advantage that no element is found in excessive amounts in the kernel.
It should be noted, however, that a considerable amount of minerals is lost
during processing in the manufacture of protein concentrates.
Derise et al. (1974) showed that, for three different cultivars grown
under similar conditions in the USA, the mineral element content was very
similar for a range of elements. They also showed, for the same range of
mineral elements, that roasting the ground nuts led to a rise in mineral
content as volatiles were lost (except in the case of sodium where a small
fall was measured). The range of chromium in groundnuts is 3--41 mg/kg in
the kernels. This amount is nutritionally significant, considering that a
chromium intake of 50-200 f,Lg/day has been tentatively recommended for
adults. Groundnuts also supply significant levels of copper in the diet: 100 g
of roasted kernels would supply all the daily needs of copper (Khalil and
Chughtai, 1983).
Akrida-Demertzi et al. (1985) did note that the level of lead in raw fat
extracted groundnut flour was 5.3 f,Lmg/kg for groundnuts grown in a
remote coastal area. This level rose to 27.6 f,Lg/kg in groundnuts grown in
an inland area of Greece noted for higher levels of air pollution (from
TABLE 6.7 Total ash content of groundnuts and groundnut products (gllOO g
DM)
Kernels Roasted Partially Defatted Protein Isolate

defatted flour concentrate
flour
Ash 1.8-3.1 1.7-8.9 4.2-5.3 3.0-4.8 2.1 0.5

Silica free 0.1-0.4
ash
Based on data from: Hoffpauir, 1953; Ladell and Phillips, 1959; Howe et al., 1965; Rao et al.,
1965a; Sekhon et al., 1970; Ayres et al., 1974; Derise et al., 1974; Quinn et al., 1975;
Conkerten and Ory, 1976; Bookwater et al., 1979; Natarajan, 1980; Alid et al., 1981; Abdel
Rahman, 1982a; Woodroof. 1983; Khalil and Chunghtai, 1983; Wallerstein et al., 1989.
TABLE 6.8 Mineral content of groundnuts and groundnut products
(mg/IOO g DM)
Raw Roasted Defatted flour Isolate Butter
Ca 44.0-87.8 55.2-91.0 92-200 40-100 46.7

Mg 157.0-200.0 174.3-196.0 10-343 10-70 148
P 137.0-470.3 288.2-538.2 80-100 188
Na 5.8-66.0 4.2-8.5 1.4--33.3 0.3--0.5 388
K 618.2-890.0 643.5-734.1 1150-1450 5-2430 682
Fe 1.2-2.0 1.3-2.1 1.5-10.9 1.9-6.7 1.6
Cu 0.74--1.60 1.3-1.7 0.6--5.2 2.1-14.6 0.7
Zn 3.3-6.2 4.4--6.7 4.7-7.0 3.0-18.2 2.9
Mn 1.7-19.0 1.9-2.2 3.1-6.1 1.6--3.4 1.4
CI 5.8-10.1 5.7-6.4 5.8
Cr 0.2-0.4 0.3--0.5 0.26
Co 0.11
AI 2.3
Mo 0.07
B 1.8-3.2
Based on data from Oke, 1967; Derise et al., 1974; Galvaro et al., 1976; Morris et al., 1978;
Lusas, 1979; Khalil and Chughtai, 1983; Salazar and Young, 1984a,b.
leaded petrol). The lead levels fell only slightly on roasting. It is interesting
to note that, although some minerals are extracted into the groundnut oil
fraction, the levels are quite low except for cobalt and lead which appear to
be concentrated into the oil fraction (Akrida-Demertzi et al., 1985). The
boron content of raw groundnuts ranges from 1.8 to 3.2 mg/lOO g and as
boron deficiency has been associated with cotyledon damage (hollow
heart) (Salazar and Young, 1984a) it is unlikely to fall below this range. It
is interesting to note that boron has also been found to modify the relative
concentration of the basic volatile aroma constituents of roasted ground-
nuts (Salazar and Young, 1984a).
6.8 OTHER COMPONENTS
A number of compounds can affect the typical groundnut flavour, nutti-

ness, sweetness and bitterness. Many of these can be altered by variety,
growing conditions, methods of harvesting and storing and processing. The
bitter flavour in groundnuts is due in part to the saponins, which are found
principally in the embryo but also in the cotyledons under certain field
conditions (Dieckert and Morris, 1958). Four different saponins could be
separated using glass paper chromatography. Maga and Lorenz (1974)
report that the total amount of free phenolic acids in defatted groundnut
Other components 203
flour was 267 mg/kg. Of the 27 different phenolic acids identified, four
were found to occur in amounts which could contribute to the astringency
of the flour.
The red skins, which represent 2.0-3.5% of the kernels, contain tannins
and related pigments that cause an undesirable colour in the protein unless
removed during initial processing. Stansbury et al. (1950) found that the
red skins contained about 7% tannin along with thiamine. These tannins
may also be responsible for some of the bitterness associated with raw
groundnuts.
6.8.1 Antinutritive factors

Many legumes contain a range of toxic factors which can adversely affect
the nutritive value of the seed. These toxic factors are a group of unrelated
chemical compounds with varying effects on metabolic processes. A con-
sistent feature is that the effect of all these factors can be reduced by
traditional processing and cooking methods and they are not considered to
affect seriously the utilization of cooked groundnut seeds. The effect they
have when consumed as raw seeds are probably minor when compared
with other legume seeds.
An example of a toxic factor which is found in insignificant amounts in
groundnuts is the report by Oke (1967) that oxalic acid occurs in Nigerian
groundnuts at a level of 4.0 g/kg. This is comparable to many other grain
legume seeds but is low when compared with some other fruits and
vegetables.
6.8.2 Trypsin inhibitors

It is widely reported that raw legume seeds contain trypsin inhibitors and
groundnuts are no exception. Tur-Sinai et al. (1972) extracted an inhibitor
from defatted groundnuts and showed that it had activity towards trypsin
and chymotrypsin (10480 TIU and 4793 CTIU respectively in 370 g of
defatted groundnut flour). They went on to show that the inhibitor lacked
the amino acids tryptophan, methionine, isoleucine and cysteine (i.e. it
contained no sulphur bridges). They suggested that the inhibitor which
they had extracted from raw groundnuts was different to that observed in
soya beans. Soya bean inhibitors have a double headed complex which can
bind trypsin and chymotrypsin at each end; in contrast, groundnut inhibi-
tor active sites are very close to each other and may even coincide. The
overall effect of trypsin inhibitors is to cause a general delay in protein
digestion which lowers the overall digestibility of the ground nut protein.
Moist heat treatments are the most effective way of reducing their activity.
Perkins and Toledo (1982) showed that almost complete destruction could
be achieved in 45 minutes at 120 DC; at 100 DC, 98% destruction was
only achieved after cooking for 180 minutes. These results confirm the
204 The composition and nutritive value of ground nut kernels
observations of Tur-Sinai et ai. (1972) that the trypsin inhibitor was stable
when boiled for 15 minutes at pH 8.
Mir and Hill (1979) showed that groundnut meals prepared by extracting
the fat using petroleum ether at 38-46 °C for 48 hours contained trypsin
inhibitor levels which were about 11 % of those found in similarly treated
soya bean meal.
Sitren et ai. (1985) showed that the trypsin inhibitor activity of raw
groundnut flour was higher than a similarly processed soya bean flour
(144.7 vs 112.1 f.Lg lectin/g flour); moist heat (121°C 15psi for 20 minutes)
was most effective at reducing the effect of the trypsin inhibitor. Dry heat
(177 °C for 20 minutes) resulted in only 76% (groundnut) and 61 % (soya
bean) inhibition of the trypsin inhibitor. In a subsequent feeding experi-
ment it was interesting to note that, after 4 weeks, the rats eating the raw
and dry-heated groundnut-based diets (which would contain high levels of
trypsin inhibitor) were consuming as much diet as the rats consuming the
moist heated groundnut diet. The rats fed the raw and dry heated soya
bean diet had considerably reduced intakes when compared to the moist
heat treatment diet. This experiment suggests that rats at least can adapt to
the trypsin inhibitor found in groundnuts and also demonstrates that it is
not readily destroyed during heat processing.
6.8.3 Haemagglutinins
A non-specific haemagglutinin has been isolated in skin-free groundnut
kernels by Dechary et ai. (1970), who noted that the amino acid compo-
sition of purified groundnut haemagglutinin was very similar to the non-
specific haemagglutinin concanavalin. The low levels of haemagglutinins
found in groundnuts and the fact that they are readily inactivated by heat
suggest that haemagglutinins are not a significant problem in groundnuts.
6.8.4 Goitrogens
Groundnuts have been reported to produce goitrogenic effects when fed at
high levels to animals (Srinivasan et ai., 1957). A phenolic glycoside is
thought to be responsible and is reported to be present in the testa, which
is commonly removed in most cooking processes. It was noted in these
experiments that the goiterogenic effect was counteracted by the addition
of iodine in the experimental diets. In iodine deficient areas, goitrogens
may have an effect, when large amounts of groundnuts are consumed in
the diet but to date no details of adverse effects have been published.
6.9 CONCLUSIONS
Traditionally, groundnuts have been considered as a source of high quality

vegetable oil; however, the annual production of protein has reached very
References 205
significant proportions. The ground nut is now the world's fourth most
important source of edible oil and third most important source of vegetable
protein. It makes a significant contribution to the diets of many people in
developing nations. The manufacture of many food products consisting of
various cereals fortified with a concentrated protein source from ground-
nuts has done much to alleviate malnutrition in many countries. The range
of products that can be made from groundnuts is perhaps the reason why
the consumption of this product has increased around the world. In part
the special taste and aroma that can be achieved on cooking, especially
when roasted, is an important feature of this food's acceptance in an
extremely wide range of cultures.
Groundnuts characteristically contain high levels of oil and protein and
low levels of carbohydrates and ash. In other ways groundnuts are similar
to many other grain legumes. Groundnut protein is deficient in some
essential amino acids (methionine and lysine) but overall the digestibility
of the protein is superior to many other plant protein sources. Groundnuts
do contain a number of anti nutritive factors but these tend to be found at
lower levels and appear to be less active than in other grain legumes. All of
these factors can be reduced by traditional processing and cooking
methods and they are not considered to affect seriously the utilization of
nutrients in cooked groundnut seeds.
Considerable potential still exists to improve the nutritive value of
groundnuts by plant breeding and by developing improved processing
techniques. The potential of groundnuts to produce large amounts of
protein should be encouraged as this vegetable protein production is very
efficient when compared with alternative animal production systems.
ACKNOWLEDGEMENTS
The assistance of the staff of the Lincoln University Library to obtain many
of the references used in this work is gratefully acknowledged.
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Journal of Food Science, 32, 129-130.
CHAPTER 7
Mineral nutrition
G.J. Gascho and]. G. Davis
7.1 INTRODUCTION
High yields of quality groundnuts require good nutrition. Sixteen elements

[(carbon (C), hydrogen (H), oxygen (0), nitrogen (N), phosphorus (P),
potassium (K), calcium (Ca), magnesium (Mg), sulphur (S), zinc (Zn),
manganese (Mn), iron (Fe), copper (Cu), boron (B), molybdenum (Mo),
and chlorine (CI)] are considered essential for plants although not all have
been proved essential for any particular species. Of the sixteen, C, Hand
o are supplied in the atmosphere or in the air space of the soil and are
called non-mineral nutrients. They are usually assumed to be plentiful,
even though 0 may be limiting due to excessive water in soil pores.
Therefore neither fertilizer nor soil amendments are applied for the ex-
pressed purpose of supplying these three elements. In addition, nickel (Ni)
and cobalt (Co) are essential for some legumes and sodium (Na), silicon
(Si), selenium (Se), and aluminum (AI) have been shown to be beneficial
in some cases. Quantities of nutrients removed by groundnut pods and
vines are presented in Table 7.1.
The groundnut is unique in one aspect of its nutrition: the seed develops
via nutrients it gathers directly from the soil rather than those transported
from roots to shoots and back to the seeds. This unique aspect has required
TABLE 7.1 Nutrient uptake/removal in groundnuts (kg/ha) (Gascho, 1992)
Plant part Yield N P K Ca Mg S
Pods 3 t/ha 120 11 18 13 9 7

Vines 5 tlha 72 11 48 64 16 8
Total 192 22 66 77 25 15
1994 by Chapman & Hall. London. ISBN 0 412 408201.
Calcium (Ca) 215
much research and it guides the applications of nutrients, especially cal-
cium, for greatest yield, quality and seed germination.
Groundnut mineral nutrition research has been carried out for several
decades (Harris, 1959). Previous reviews of the subject have covered the
available knowledge up to 1982 (York and Colwell, 1951; Gillier and
Silvestre, 1969; Reid and Cox, 1973; Cox et al., 1982). However, in the
decade following the last of these reviews, additional advances have been
made in our understanding of mineral nutrition of the groundnut. This
understanding has been applied to enhance yield and quality of the crop. In
this chapter, emphasis will be placed on important principles of nutrition
and particularly to advances made in the past decade. Each required
element is presented individually.
7.2 CALCIUM (Ca)
Calcium is often considered the essential element most commonly deficient

for groundnut in non-calcareous soils. The consequences of Ca deficiency
are blackened plumules, high incidences of pod rot and unfilled pods
(termed 'pops') resulting in low yield and substandard grade. Groundnuts
produced in Ca deficient conditions will exhibit much poorer germination
than those produced with an adequate Ca supply. The effects of poor Ca
nutrition are elaborated on later.
7.2.1 Needs related to uptake

For the groundnut, the exceptional requirement for Ca is, for the most
part, not in order to grow a healthy plant but for developing a properly
filled pod with a high quality seed. Bell et al. (1989) evaluated the external
Ca requirements of six tropical legumes in flowing solution culture and
found that the Ca requirements of red spanish groundnuts for growth and
nodulation were not great in relation to pigeon pea, guar, soybean and
cowpea. Guar and pigeon pea required a concentration of> 50 f..LM Ca for
nodule formation while groundnuts had nodules at 2 f..LM Ca. Maximum
growth of groundnut was found at 50 f..LM, soybean at 100 f..LM and cowpea
at 2500 f..LM Ca. Even though Ca requirements for growth and nodulation
of the groundnut are not great, the exceptional needs of Ca for groundnut
seed maturation and quality are stressed below.
Barber (1984) stated:
Calcium appears to have its greatest significance in providing the
appropriate balance for levels of other nutrients within the plant.
Dependent on the ambient Ca concentration near the absorbing
surface mass flow and diffusion can be the dominant mechanisms of
Ca absorption by roots.
216 Mineral nutrition
For most plants, Ca is oversupplied to the root surface by mass flow and
only a small proportion of the Ca is absorbed and subsequently translo-
cated via the xylem to leaves in response to the flow of the transpiration
stream. Much of the Ca is deposited in the leaf as Ca oxalate and then it is
not re-translocated. This is a different mechanism from that employed in
root uptake of several other ions. In the root, the uptake of many ions is
termed 'active', indicating that the roots select the required nutrient ele-
ments from the soil solution. Uptake of Ca is fairly 'passive', i.e. roots take
up the elements as they are presented in the soil solution without a
selection mechanism. Therefore, the amount of Ca taken up by the plant is
dependent both on its concentration in the soil solution and on the amount
of water moving into the plant (Mengel and Kirkby, 1982). Because Ca is
not easily translocated within the plant, Ca needed during reproductive
growth must be directly supplied by uptake.
Classic research with the groundnut has shown that Ca is transported
upward in the plant from root uptake via the xylem, but little or no Ca is
transported from the leaves downward through the pegs to the developing
pod via the phloem (Bledsoe et at., 1949; Wiersum, 1951; Mizuno, 1959).
The Ca absorption problem for the developing fruit is compounded by the
fact that there is little water movement upward from the peg to the plant
tops in order to provide a gradient to carry needed Ca into the developing
fruit via mass flow (Wiersum, 1951; Beringer and Taha, 1976; Wolt and
Adams, 1979).
Calcium for seed development must be absorbed by the peg via passive
uptake by diffusion (Sumner et at., 1988). Therefore, it is clear that
a relatively high concentration of Ca is needed in soil in the pod
development zone in order to avert Ca deficiency. Calcium in that
zone must be replenished throughout peg and seed development
periods in order to maintain a gradient of Ca2 + toward and into the pod.
The developmental period extends from 20-80 days following the entrance
of the peg into the soil. Mizuno (1959) determined that 92% of the Ca was
taken up by the pod during that time period and 69% was taken up
between day 20 and day 30. Smal et at. (1989) found that withholding Ca
from the pod zone during the first 30 days following initial pegging resulted
in the smallest seeds and the least seed dry weight. They determined that a
continuous Ca supply of 3.75 f.LM Ca solution resulted in greatest seed size
and weight, but suggested that the 30-day period following pegging is most
critical.
For soils with great Ca supplying power and rapid replenishment of soil
solution Ca, no deficiency is expected. However, the ground nut is often
grown on sandy soils which possess limited ability to supply Ca to replenish
the soil solution. This problem is compounded because such soils are often
relatively droughty and the groundnut, being relatively drought-tolerant in
comparison with most plants grown in semi-humid regions, is often grown
in localities with limited rainfall during some portion of the period of peg
Calcium (Ca) 217
and seed development. Limited soil moisture at this time worsens the
situation in droughty soils: added Ca compounds will not dissolve without
moisture, so that there is a lack of Ca solution; there is also a lack of a
diffusion gradient so that Ca2 + cannot move toward the pod. Therefore,
Ca deficiency results.
Effects of drought on Ca uptake by developing fruits have been reviewed
by Boote et ai (1982). They emphasized studies showing poor uptake in
drought periods and the enhancement of uptake with adequate water in the
fruiting zone. Groundnuts showing Ca deficiency symptoms during periods
of drought also had lower Ca concentrations in seeds and hulls than those
showing no Ca deficiency symptoms. Alva et ai. (1991) determined soil
solution Ca following gypsum application and incubation in the laboratory.
They imposed four drying cycles over a period of 70 days in two soils that
represent the textural extremes of the production industry in Georgia.
Following the drying cycles, the soils were returned to field capacity
moisture and soil solution Ca was determined. Soil solution Ca increased
with increasing soil moisture in both soils for the first 14 days following
gypsum application. Subsequently, soil solution Ca decreased for all
moisture regimes in the Bonifay sand (grossarenic Plinthic Kandiudult),
but this was true only in the driest regime for the Greenville sandy loam
(thermic Rhodic Kandiudult). The results indicate that soil solution Ca
deficiency is especially likely in soils such as the Bonifay sand due to low
moisture retention. Calcium nutrition for the developing nut is intimately
associated with soil moisture. For this reason it appears simplistic to cite a
critical soil Ca test level for groundnuts. The true 'critical level' will depend
on the moisture during nut development and will change, both geographi-
cally and temporally, even for a given cultivar.
Concentrations of other cations in the soil, particularly K and Mg, can
affect Ca uptake and thereby affect yield and quality. Several reports of the
deleterious effects of high soil K and seed K on groundnuts have been
published (Bolhuis and Stubbs, 1955; Hallock and Allison, 1980; Alva et
ai., 1989a, 1991; Lynd and Ansman, 1989). Top growth is depressed with
the addition of K alone, but increased by the addition of K + Ca (Lynd and
Ansman, 1989). Using the Mehlich 1 (double acid) extractant, Alva et ai.
(1989a) determined that an optimum CalK ratio in the pegging zone topsoil
is near 10: 1.
7.2.2 Genotypes and Ca nutrition

In general, greater soil concentrations of Ca are required for larger seeded
than for smaller seeded cultivars (Walker et ai., 1976; Walker and Keisling,
1978). However, Cox et ai. (1982) cited several exceptions. Gaines et al.
(1989) determined both runner (cv. Florunner) and virginia type (cv. NC-7
and Early Bunch) responses to gypsum application relative to Mehlich 1
soil Ca. Yield response in the Florunners was limited to experiments with
125,-----~--~----~----~----~--~----~--__.
";" : .
l 100 .........: •. r/~' ~:E::;~1·~:.r~;~~J':···~;·'·······";·············
"0
75 ...... ;,..... /:. ::.:.<.. j.....•...... ;.~ ..............j..............~ ............. :............. j............. .
Qi
'>,
OJ
>
~
/<}/. i ' i
50 ....... ,.....,....... ~' ,
"f
,
!
,
,
,
i,
Qi
"IrIT!:
a:
25
ii
o
o 200 400 600 800
Mehlich 1 Ca (mg/kg)
Figure 7.1 Relative pod yield of virginia type receiving no calcium additions to
those receiving calcium as affected by Mehlich 1 extractable calcium in soil prior to
addition of any calcium. Dotted lines are 95% confidence levels. Data from 122
experiments conducted in Alabama, Georgia, North Carolina and Virginia.
Mehlich 1 soil Ca less than 270 mg/kg, while the virginia type responded to
all levels of soil Ca available in the experiments (Gaines et al., 1989). Data
from experiments conducted in Alabama, Georgia, North Carolina and
Virginia have been combined to show the effect of Mehlich 1 soil Ca on
relative yield of the virginia type (Hodges et at., 1993). Maximum yield was
attained at about 525 mg Mehlich 1 Ca/kg (Figure 7.1). Likewise, experi-
ments conducted in Alabama and Georgia were combined to indicate that
maximum yield of the runner type was attained at a Mehlich 1 Ca test of
approximately 225 mg Ca/kg (Figure 7.2).
No good evidence exists to show that the internal Ca requirements of
groundnuts are affected by seed size. Adams et ai. (1993) found that the Ca
concentrations in seed which produced maximum germination ranged from
381 to 414 mg/kg for four small-seeded runner cultivars. Those values are
close to the 420 mg/kg which was found to give maximum germination of
Florigiant, a larger-seeded cultivar, in an earlier study in North Carolina
(Cox et al., 1976). Gascho et al. (1992) correlated germination with seed Ca
to find that the large-seeded virginia type (cv. GK-3) had an internal Ca
requirement of approximately 600 mglkg, which is similar to the internal
Calcium (Ca) 219
requirement for maximum germination found for the small-seeded runner
type (cv. Florunner) and considerably greater than those reported by
Adams et at. (1993) or Cox et al. (1976). At least for the present, the need
for greater ambient Ca concentrations in soil for the large-seeded cultivars
is explained by acceptance of the view that diffusion from soil solution is
the mechanism by which Ca enters the pod. Sumner et at. (1988) evaluated
eight genotypes with widely varying pod and maturity characteristics over
two seasons and four drought stress periods applied at different times
during the crop and found strong relationships between Ca concentration
in the hull and surface area of the pod. A larger-seeded pod has a lesser
ratio of surface area to weight than a smaller-seeded pod, so that it is less
efficient in diffusion and requires a greater concentration of soil solution
Ca and/or greater soil moisture in order to provide adequate Ca to the pod.
Acceptance of diffusion as the mechanism by which Ca moves to the
developing peg explains the observed need for greater Ca applications for
the large-seeded cultivars.
Variables other than the ratio of seed size to surface are involved in
external Ca needs among cultivars. Kvien et at. (1988) used stepwise
regression to select five genotypic characteristics which significantly influ-
ence seed and hull Ca concentrations:
125,-------~--~--------------~----------~--__.
"
""-».... .!.,:.....-r-'-\'--....;!--:---+--.-----:-+-----+-----j
-:. : ::. . • 1: ••
100 ........•.•• ';..., - " .. :. -.0-'."'.
75
:~ /~.i.:;"~:'~:";L-":" ".i.,-'...........
. . _ _ ". ~ :'
/i/- j.
;,
j
:
j
:
j
:
j
:
j
~
:.1' ..............,_____________ .'- ____________ _____________ _
:
j
'i" " , i' i : i i i

50 ....,i i ...... :............. ~.····· ........ : ............. ~ ..............: ............. : ............. ~ ............. .
" ! ' , ! ' !
25 "·:·······················;TT'T
,,, , ,,, ..
, ., , ,
OT-----,,-----r-----+'----~'------'r-----r'-----+---~
o 200 400 600 800
Mehlich 1 Ca (mg/kg)
Figure 7.2 Relative pod yield of runner type receiving no calcium additions to
those receiving calcium as affected by Mehlich 1 extractable calcium in soil prior to
addition of any calcium. Dotted lines are 95% confidence levels. Data from 168
experiments conducted in Alabama and Georgia.
• days required to mature a pod;
• specific hull weight;
• pod surface area;
• hull thickness;
• pod volume.
Cox et al. (1982) suggest that Ca efficiency may be increased by breeding
the Ca-efficient characteristics of small-seeded runner types into the large-
seeded cultivars. Recent literature does not reveal much effort in this
regard. In an attempt to identify groundnuts suitable for culture on
marginal soils and for low-input agriculture, Branch and Gascho (1985)
evaluated 24 cultivars of four USA market types (runner, virginia, spanish
and valencia) for their tolerance of low soil fertility. They found that the
standard south-eastern USA runner-type cultivar, Florunner, gave the
greatest pod and total sound mature kernel yield but the kernels had a
great incidence of concealed damage, indicative of Ca deficiency. The pre-
eminence of the Florunner cultivar may be due, at least in part, to its Ca
efficiency - defined as its ability to yield in a low ambient Ca environment.
However, improvement of nutrient efficiency does not appear to be a
major objective with breeders in their efforts to improve yield, grade and
disease resistance and to decrease aflatoxin concentrations. Recent
evaluations of the effects of supplemental Ca application on yield, grade
and seed quality of promising runner-type cultivars (Adams et al., 1993)
suggest that these cultivars may require different ambient soil solution
Ca from that for Florunner. In their study, Ca concentrations in seeds
averaged by cultivar across 14 sites and for plots receiving and not
receiving gypsum were 360 mg/kg for Florunner, 356 mg/kg for Sunrunner,
345 mg/kg for GK-7 and 301 mg/kg for Southern Runner. These results
indicate that no promising runner-type cultivar is more efficient than
Florunner in Ca absorption. The relatively new Southern Runner cultivar
was less efficient, especially in cases where no gypsum was applied: seed Ca
concentration for Florunner was 319 mg/kg, and for Southern Runner
235 mg/kg.
7.2.3 Symptoms of Ca deficiency

The important consequences of Ca deficiency occur in the reproductive
stages of development, but some indication of insufficiency for quality seed
production may be evident in the vegetative stages of growth. Symptoms
summarized by Cox et al. (1982) from the literature include:
• more abundant foliage that remains green later in the season;

• tendency for a greater number of flowers, many of which may be
infertile;
Calcium (Ca) 221
• localized pitted areas on the lower surface of fully developed leaves
that can later develop into brown chlorotic spots, which may have halos
on their perimeters, coalesce and cause senescence of the leaves;
• death of root tips and terminal buds (only in extreme deficiency).
Death of seedlings due to Ca deficiency occurred in a sandy field when
the Mehlich 1 soil Ca level in the upper 15 cm was 6-21 mg/kg (Gascho
et al., unpublished data). Pod production, grade, germination of the seed
produced and seedling survival were reduced greatly by and are the final
results of Ca deficiency. The primary reason for these results is the high
proportion of pods with aborted and shrivelled fruits ('pops ') and darkened
plumules ('black heart'). These deleterious effects are strongly related to
concentrations of Ca in the soil as well as in seed and hull (Adams and
Hartzog, 1980; Hallock and Allison, 1980; Walker and Csinos, 1980).
Germination of seed produced in low Ca soils is greatly reduced. Harris
and Brolmann (1966c) determined 100% germination and 98% seedling
survival when seed Ca was >800 mg/kg and 23% germination with no
seedling survival for seeds with <300 mg/kg.
Garren (1964), Hallock and Garren (1968) and Walker and Csinos
(1980) provide data indicating that pod rot (caused by several organisms,
but especially Pythium myriotylum Drechs. and Rhizoctonia solani
Kuehn.) is related to low Ca supply, based on experiments where pod rot
incidence was reduced by application of gypsum. Hallock and Garren
(1968) and Csinos and Gaines (1986) suggested that the balance among Ca,
K and Mg was important, with high K or Mg supply increasing the
incidence of pod rot. The reported effect of Ca on pod rot has been
disputed by Filonow et al. (1988), who infected soil with Pythium myrioty-
lum Drechs. and Rhizoctonia solani Kuehn. and found that increasing Ca
applications had no effect on pod rot incidence. Their work supports the
effect of the organisms on the aetiology of pod rot, but argues against Ca
deficiency in the hull as a primary cause of the problem. Recent work in
Georgia supports the relationship between Ca and pod rot. Gascho et al.
(1993) conducted field experiments in soils with a Mehlich 1 soil Ca of less
than 200 mg/kg and found that pod rot averaged 8-9% in control plots of
both the small-seeded Florunner and the larger-seeded GK-3. Pod rot
incidence was decreased significantly by the application of Ca.
Calcium supply may also be related to aflatoxin infection. Davidson et al.
(1983) and Wilson et al. (1989) determined that Ca applications on sandy
soils with a low Ca status decreased infection by Aspergillus fiavus, which is
sometimes associated with high levels of aflatoxin production.
7.2.4 Diagnosis of Ca needs

Soil analysis is the most useful diagnostic tool for determining the need for
supplemental Ca fertilization. Samples taken from the surface 15-20 cm
prior to planting the crop are often used. The choice of critical Ca level
varies with the soil extractant and the type of groundnut grown. Adams
and Hartzog (1980) determined that 125 mg/kg of Mehlich 1 extractable Ca
was the critical value for cv. F10runner in Alabama. Hodges et al. (1993)
consolidated data from the literature for 168 experiments conducted in
Alabama and Georgia and determined that 95% of maximum yield was
obtained when the Mehlich 1 extractable Ca was 200 mg/kg (Figure 7.2). In
practice, supplemental Ca applications for runner groundnuts are normally
recommended for soil tests less than 150 (Alabama) to 250 mg Calkg
(Georgia) to help ensure adequate Ca (Cope et al., 1981; Plank, 1989a).
For the larger-seeded virginia type, soil test Ca is less valuable since
supplemental Ca is normally recommended regardless of the soil test level
(Plank, 1989a). Gaines et al. (1989) indicated 95% of maximum yield was
attained for virginia type at a Mehlich 1 test reading of approximately
775 mg Ca/kg. Parker et al. (unpublished) combined data from the litera-
ture for 88 experiments conducted with large-seeded virginia types in
Alabama, Georgia, North Carolina and Virginia. They determined that
98% of maximum yield was obtained when Mehlich 1 Ca was 525 mg/kg.
No doubt there is a soil Ca level above which no yield or quality responses
to applied Ca would be expected for large-seeded cultivars but some
additional data, particularly in relation to grade, is needed before revising
present gypsum recommendations for the larger groundnut. For all types, a
particular soil test value should not be chosen as a 'critical level', as that
terminology implies precision greater than possible in farmers' fields. The
level of soil test above which a response is not expected will vary with many
genotypic characteristics but also due to the soil moisture available to
transport Ca to the developing fruit.
An ideal soil test would evaluate Ca concentration in the soil solution
just prior to bloom. Applications of a slowly soluble Ca source (such as
gypsum) could be made to supply Ca if needed. Attempts have been made
to measure concentrations directly in the soil solution or to use weak
extract ants that mimic the soil solution without solubilization of normally
soil-solution-insoluble Ca. Smal et al. (1988) evaluated 0.01 N NaN0 3 in
comparison with a stronger extractant (Mehlich 1) which is widely used to
measure cations in the acid soils of the south-eastern USA. They con-
cluded that yield and grade parameters were better related to Ca levels
produced by the NaN0 3 extractant than Ca by the Mehlich 1 solvent. Such
tests with weak extract ants should be"valuable for measuring soil Ca in the
pegging zone, especially following the application of limestone prior to
planting as they should not solubilize limestone particles to the same
degree as the stronger extractant.
However, later studies have re-emphasized that Mehlich 1 is the superior
extractant (Alva et al., 1991). However, it may give unreasonably high Ca
levels in cases where limestone has been applied recently and where
sampling may encounter limestone particles which can be dissolved by
Calcium (Ca) 223
Mehlich 1. In some cases, Ca extracted by NaN0 3 was less correlated with
peanut Ca needs than was the standard extractant for soil Ca (Mehlich 1) in
experiments conducted in Georgia. Alva et al. (1991) found that the NaN0 3
extractant did not predict a need for supplemental Ca on a very sandy site,
but Mehlich 1 extraction accurately predicted a response. Adequate soil
solution Ca (evaluated by a weak extractant) for such a soil is no guarantee
that adequate Ca will be available when required by the groundnut. Mehlich
1 extracts are more reliable for predicting Ca needs than are 0.01 N NaN0 3
extracts on soils with low ability to hold Ca. The limitations of the weak
extract ants probably occur because such tests make no allowance for or
evaluation of the Ca which may become available during the developmental
period in which Ca is in constant demand. That time can vary with cultivar
and locality, but can be for a period of up to 60 days in the peanut belt of the
south-eastern United States. Need for supplemental Ca application to soils
which have little ability to supply Ca from reserves can be grossly underesti-
mated by soil solution Ca or by weak extractants.
If limestone is applied by the preplant incorporated method (section
7.2.5), the post-planting soil test will be of less value as there will probably
be some lime particles in the pod development zone that may be dissolved
by the strong extractant, raising the test to a very high value. However,
those particles may not be dissolved in the pod developmental period, in
time to benefit the current crop.
Foliar analysis has been used to a limited extent. Due to the unique
method of uptake of the developing peg discussed above, it is easy to
understand why it has been difficult to obtain satisfactory correlations
between leaf nutrient concentrations, yield and seed quality parameters.
Nicholaides and Cox (1970) predicted a Ca deficiency when Ca in the tops
of 9-week-old plants is less than 1.2%. Others have determined different
critical levels varying quite widely (Cox and Reid, 1964). Determination of
foliage Ca concentrations is diagnostic for gross deficiencies in growth of
the plant, but not as a tool that should be relied upon to ensure a good
harvest. Sufficiency levels in leaves are given in Table 7.2.
7.2.S Correction of Ca deficiency

Getting soluble Ca into the pegging zone and maintaining a supply of that
element to replenish depletion by the crop and losses to leaching can be
especially difficult in a sandy soil with a poor Ca supply. Very soluble Ca
sources such as CaCI2 are generally expensive and short-lived due to
leaching losses. Gypsum (calcium sulphate) is much less soluble, but its
application on the soil surface at first bloom is generally appropriate for
supplying Ca over the 45-60 day period in which the peanut has a great
requirement (Sridhar et al. 1985).
In addition to mined gypsum, by-product and phosphogypsum (Alva
et al. 1989b, Gascho and Alva, 1990) are available in many places at lower
cost. When soil Ca is less than the chosen threshold level, gypsum is either
broadcast or banded over the groundnut row at first bloom. Most products
have a range of particle sizes that solubilize over time to provide Ca.
Gypsum has been a very successful source of Ca; however, its application
can add considerably to production costs. Dependent on source and
recommended rate of application, the applied costs may range from about
$40 to $80/ha.
Limestone is an important source of Ca for crops grown in the acidic
soils of south-eastern USA and other locales. The main response to
limestone is due to its supply of Ca, whether applied for that purpose or to
increase soil pH. Adams and Hartzog (1980) conducted 16 field experi-
ments with runner-type groundnuts in Alabama with the conclusion that:
'Lime appeared to do little more than serve as a source of calcium.
Spring-applied lime provided all of the Ca needed for maximum yield and
grade when it was properly incorporated into the pegging zone.' Since
good farmers will lime their soils for their chosen crop rotation, they
should attempt to apply it in a manner that will supply Ca to their crop and
thereby reduce or eliminate the expense of a gypsum application. Adams
and Hartzog (1980) have shown that, when limestone is applied to the soil
prior to planting and incorporated to a shallow depth of 5-12 em, adequate
TABLE 7.2 Sufficiency levels of nutrients in groundnut leaf dry matter (Gillier and
Silvestre 1969; Plank, 1989b)
(a) Macronutrients (%)
Plant Time N P K Ca Mg s
part
7th leaf 40 days 3.3-3.9 0.15-D.25 1.0-1.5 2.0 0.3 0.19-D.25

after
planting
Upper Bloom 3.0--4.5 0.20-0.50 1.7-3.0 1.25-2.0 0.3-D.8 0.2O-D.35
mature
leaf
(b) Micronutrients (mg/kg)
Plant Time Mn Fe Zn Cu B Mo AI
part
Upper
mature
leaves Bloom 20-350 50-300 20-60 t 5-20 20-60 0.1-5.0 <200
tCa:Zn ratio <50:1.

Calcium (Ca) 225
TABLE 7.3 Effect of limestone placement on soil pH and Ca on groundnut yield
and grade
Soil
Placement pH Ca Pod yield SMK2
mg/kg kg/ha %
Turned under 5.7 32 2285 66
PPI 6.4 155 4426 73
No lime 5.7 30 2455 66
1 2.2 tons dolomite limestone/ha. either turned under with a mouldboard plough, or applied
on soil surface following turning and incorporated approximately 8 em (PPJ), or no lime
applied.
2 SMK, sound mature kernels
Ca is normally available for runner-type groundnuts as judged by the fact

that responses to gypsum applied at bloom were rare following such
applications. Gascho and Hodges (1991) confirmed these results in
Georgia, but found that limestone incorporated before planting, applied at
rates adequate to increase pH to the recommended value, did not supply
adequate Ca for the larger seeded virginia type. On the other hand, Bell
(1985a),working with Virginia Bunch on the irrigated sands of the Ord
River area of Australia, determined that shallow incorporation of 1000 kg!
ha of limestone (90% CaC03 ) before planting produced maximum yields
of sound mature kernels. In his study, additional applications of gypsum at
flowering had no significant effects on yield.
Much of the inconsistency in reported results may be due to the prevail-
ing moisture conditions. A wet growing season will result in good solution
Ca regardless of whether the Ca source is gypsum or limestone. If con-
ditions are extremely wet on a deep sand, there is the possibility that Ca
may be leached to a depth below the pegging zone. Application of gypsum
at bloom followed by dry soil conditions can result in poor solution of Ca,
while heavy rainfall on such a soil could result in Ca leaching.
Limestone placement is critical in evaluating its usefulness to the crop.
Most limestone applied in agricultural fields is incorporated much deeper
than the pegging zone either by harrows or ploughs. Nearly all groundnut
fields in the US are turned by mouldboard ploughing in order to bury trash
and provide a loose bed for nut development and removal. The modern
mouldboard plough essentially inverts the top 25-30 cm of soil. If lime-
stone is applied prior to ploughing, it will remain far below the zone of nut
development and not provide Ca needed by the groundnut. Gascho and
Hodges (1991, Table 7.3) found no benefit to runner-type cultivars in
preploughing application and benefits equal to gypsum application for
limestone incorporated before planting.
Unfortunately, the practice of shallow incorporation of limestone has
been adopted only slowly in many growing areas of the world. In some cases,
limestone is not available or is too expensive, and no Ca is applied. In other
cases, limestone is ploughed under or otherwise incorporated too deeply into
the soil or gypsum is considered the only appropriate source of Ca.
Dolomite is the limestone of choice when soils are deficient in Mg but,
for the groundnut, the soil test must be very low to derive a benefit from
Mg application. Adams and Hartzog (1980) determined that dolomite was
equal to calcite as a Ca source (consideration must be given to the lower Ca
concentration in the dolomite) for all cases tested in Alabama except one
where Mehlich 1 Mg was 4 mg/kg. Gascho and Hodges (1991) also found
dolomite to be superior, on sandy soils in Georgia, only when soil Mg
was very low. However, most rotational crops have a greater Mg require-
ment than groundnut, and their needs must also be considered in a liming
programme.
7.3 NITROGEN (N)
Approximately 190 kg/ha of N is removed with a 3 ton/ha groundnut pod

crop when the vines are also removed (Gascho, 1992). The groundnut is a
legume and under most conditions enough N is fixed through symbiotic
relations with Bradyrhizobium spp. (Chapter 8) to avoid deficiency
throughout the plant's life-cycle, including the production of a good yield
with high quality. During reproductive stages, N is continually mobilized
from leaves to the developing fruit (Kvien et al., 1986) and under some
conditions N deficiency can occur. For that reason, some N is commonly
applied to groundnuts in most growing areas of the world (Gascho, 1992).
However, N is not recommended in the USA, except in cases where
symbiotic fixation is low or expected to be inefficient. In China, 30 kg N/ha
is commonly top-dressed following an initial application of organic manure
and PK fertilizer. In India, 10-25 kg N/ha is commonly applied; in Senegal
the recommendation is approximately 12 kg N/ha.
Conditions that affect N-supply to the groundnut include type, variety,
presence of inoculum, crop rotation, soil type, soil moisture and tempera-
ture. Cox et al. (1982) summarized N fertilization studies prior to 1982 by
stating:
There seem to be a number of conditions conducive to obtaining a
response from fertilizer N. The sequentially branched botanical
varieties are more responsive than the alternate branched. In India,
responses to N seem more likely if P is also low, but this did not hold
true for more acid soils in South America. Nitrogen response could
not be shown if other factors were more limiting. Substantial amounts
of N were not ordinarily needed unless the site was exceedingly low in
N or no effective rhizobia were present.
Nitrogen (N) 227
Studies since 1982 have not appreciably changed any of the previous
interpretations of the needs of groundnut for fertilizer N. Several studies
still lead to differing conclusions, due to variations in the factors given
above. Walker et al. (1982), Hiltbold et al (1983) and Cope et al. (1984)
found no responses to N application for the runner type in Georgia and
Alabama. In further studies, Walker et al. (1984) found that yields of non-
nodulating groundnuts and Florunner and Tifrun cultivars were linearly
increased with the number of foliar applications of N at 13.5 kg/ha begin-
ning 28 days after emergence. Patel et al. (1988) found that N application
increased yield in only one of five years of their study in India. Mali et al.
(1988) found increased N uptake and yield due to application of 20 kg/ha at
planting in India, while Lal and Saran (1988) also measured increases in
pods/plant and in oil and protein contents.
Inoculation of seed with a correct rhizobium strain seldom increases
growth and yield by alleviation of N deficiency. Cox et al. (1982)
stated:
Native rhizobia are abundant and apparently able to fix adequate N
at current yield levels. Both host plants and rhizobia strains may be
selected to improve N uptake, so a means of improvement is avail-
able if needed. Neither management practices nor most soil con-
ditions have been shown to affect N fixation greatly, but the rate may
be decreased substantially by adverse climatic conditions.
In 1983, J.e. Wynne and associates in North Carolina began publication
of several studies on N fixation. In the initial experiment, Ball et al. (1983)
found no increased plant weight or yield in either spanish or virginia types
due to inoculation with rhizobium, while application of N fertilizer
increased both plant weight and yield of both groundnut types but reduced
N-fixing rates. From their data they suggested that there is no need for
inoculation when the soil contains a high population of the proper strain of
rhizobium, but either the groundnut-rhizobia symbiosis does not fix
enough N for maximum yield or fixed N is not used as efficiently as
fertilizer N. Arrendell et al. (1985) used host selection in an attempt to
increase symbiotic N fixation and found a significant correlation of nitroge-
nase activity with shoot weight, but lower correlations between nitrogenase
and yield. They concluded that selection for greater N-fixing activity
should be possible and should result in indirect selection for improved
yield. However, in a later study, Arrendell et al. (1988) were unable to
generate any variability in N-fixing ability among F2 selections. Finally,
Phillips et al. (1989) determined the presence of non-additive genetic
effects in two crosses and concluded that early generation selection for N
fixation factors would be ineffective. Thus it appears that little progress has
been made in improving N fixation by selection to date.
Nitrogen application has been considered a control measure for cylin-
drocladium black rot (CBR) in some instances. Black et al. (1984) found
that N decreased CBR severity for nodulating groundnuts when inoculum
density was low but increased CBR severity in non-nodulating lines.
Nitrogen concentrations in leaves decrease with plant age and vary with
cultivar, making leaf N a difficult diagnostic tool for determination of need.
Analysis of the seventh leaf 40 days after planting should be in the
sufficiency range of 3.3-3.9% (Gillier and Silvestre, 1969; Plank, 1989b).
7.4 PHOSPHORUS (P)
The groundnut is grown on P deficient soils in many areas of the world.

On a global scale, P may be the most deficient element. However, the
deficiency is primarily limited to areas which have never been fertilized
with P, where P fertilizers are not available or their cost is prohibitive.
Phosphorus deficiency in groundnuts is well documented in several areas
(Cox et al., 1982; Chauhan et al., 1988; Bell, 1985b; Mali et al. 1988;
Survanvesh and Morrill, 1986; Dwivedi et al. 1987) but is unusual in
countries such as the USA, where P fertilizers are commonly used (Cope et
al. 1984; Walker et al. 1982). In general, P deficiency in groundnuts can be
corrected easily by application of P fertilizers since the crop is grown on
sandy soils with low amounts of clays - soils in which P fixation is not a
common problem. Fertilization generally not only provides enough P for
the crop but also increases available P in such soils since the plant require-
ment and removal are low (Table 7.1 ) and little P is lost by leaching, even
in sandy soils. An exception is made for groundnut culture on calcareous
sands, where P fixation can lead to P deficiency in spite of fertilization.
Soil analysis is considered the best method to determine P fertilizer
needs. Effectiveness of extract ants varies between locations and care must
be exercised in comparisons of 'available soil P' values in the literature.
The predominant extractants, proposed critical levels for groundnut and
responses obtained in groundnut soils throughout the world are discussed
by Cox et al. (1982). These will not be discussed here.
Soil P levels required for groundnut are often lower than those required
for other crops (Cope et al. 1984). A summary of data from India, Bolivia,
South Africa, Guyana, Colombia, Israel, Uganda, China, Australia and
the USA, with a number of extractants, suggest very low critical levels of
approximately 10 mg P/kg. A review of recent research emphasizes the low
P requirements of runner varieties. In Alabama, Hartzog and Adams
(1988a,b) found no correlation between Mehlich 1 extractable P and either
yield or grade of runner groundnuts from fertilization in 39 experimental
sites where soil P ranged from 1 to 45 mg/kg. Cope et al. (1984) found no
response to P fertilizer in long term experiments even when Mehlich 1 P
was reduced to 11 mg/kg by continuous cropping without fertilization. The
critical level for Mehlich 1 P in the south-eastern runner-type peanut belt is
approximately 10 mg/kg (Mitchell and Adams, 1993).
Phosphorus (P) 229
Bell (1985b) found responses to applied P at very low levels of 0.5 M
NaHC0 3 extractable P on sands in Australia. He determined that the
critical level for NaHC0 3 soil P was 7.3-7.9 mg/kg for Virginia Bunch.
Chauhan et ai. (1988) also had some responses, but only at very low levels
of soil P (6.15 mg/kg, unknown extractant). Since 1982, the Mehlich 3
extractant has been introduced and is now in use in North Carolina and is
under consideration in other south-eastern states currently using the
Mehlich 1 extractant (formerly called Double Acid). Mehlich 3 commonly
extracts 1.5 to 2 times more P than Mehlich 1 (Gascho et ai., 1990).
Therefore critical levels are considered to be proportionally greater for
Mehlich 3. The corresponding critical level for Mehlich 3 is 20-24 mgldm 3
(Cox, unpublished data). In areas where P fertilizers are readily available
and relatively inexpensive, maintenance of such low levels is nearly assured
by P applications to rotational crops, and P fertilization of the groundnut
crop is generally not needed.
For areas where P fertilizers are not plentiful, or where P is fixed due to
highly calcareous conditions and responses to P have been noted, some
lines may be developed which are tolerant of low soil P. Dwivedi et ai.
(1987) conducted field experiments on a P deficient calcareous soil and
noted variation in dry matter accumulation and P accumulation in plant
parts for one adapted line in comparison with five unadapted lines. From
their results, they proposed a model for high-yielding P deficiency resistant
plant type: R = (Ay x 1.32) x (An X 1.09), where R is the resistance
value for P deficiency or low P supply, A is an adapted cultivar, y is yield, n
is P uptake on a P deficient soil, and 1.32 and 1.09 are constants. The
authors claim that the model will predict the resistance of lines or cultivars
to P deficiency.
Some researchers have considered foliar applications of P in cases where
soil applications are fixed by great concentrations of Fe, AI, or free phase
CaC03 . Survanvesh and Morrill (1986) made foliar applications of P in the
greenhouse to find that it was effective in increasing growth and plant P
concentrations when the supply of P to the roots was inadequate, but not
when root availability was adequate. In field experiments, Walker et ai.
(1982) found no response to foliar P when adequate preplant fertilizer was
applied on a non-fixing soil.
Plant P analysis has also been useful, to an extent limited by the fact that
effective applications of P cannot usually be made following a diagnostic
tissue test. Therefore plant analysis is only helpful in knowing what to do
when groundnuts are next planted in a given field. Care must be taken in
the interpretation of plant analysis results since concentrations vary signifi-
cantly and by plant part and age. Bhan (1977) noted that leaf P declined
from 0.35% to 0.15% between 30 and 120 days following planting.
Fertilization with P, which increased yields, only increased P concentration
by about 0.05% throughout that period. Partitioning of uptake indicates
that only 10% of the total is taken up in the vegetative phase while 39%
and 51 % are taken up in the reproductive and maturation stages, respect-
ively (Longanathan and Krishamoorthy, 1977). Foster (1980) set a critical
P level of 0.29% P for leaves at flowering. Others use a range of sufficiency
levels and a compilation of these (Gascho, 1992; Gillier and Silvestre,
1969; Plank, 1989b) indicates a wide variance. Reported ranges are 0.15-
0.25% for the seventh leaf 40 days after planting and 0.25-0.50% for upper
mature leaves at bloom.
7.5 POTASSIUM (K)
Removal of K from soil by groundnuts is considered to be low relative to

soybeans or bermudagrass when the vines are returned (Table 7.1).
Removal of the vines as hay quadruples K removal. These data should be
considered in planning a cropping system but may not be of great import-
ance when it is grown in a rotation with several other crops, since ground-
nuts will not respond to direct K fertilization unless soil available K is low
(Scarsbrook and Cope, 1956; Walker et al., 1979). Reports ofresponses to
K application are not consistent.
Scarsbrook and Cope (1956) reported an average pod yield response of
170 kg/ha to K fertilization when the Mehlich 1 soil K test was rated 'low'
by the soil K interpretation in Alabama. They found no response when K
was rated 'high'.
However, the world literature reports rather mixed responses to applied
K and, in many cases, does not provide good supporting evidence or
information on soil K levels. Piggot (1960) established that K application
was necessary for groundnuts in Sierra Leone. Potassium is rarely needed
in India but is added as insurance (Kernick, 1961). Goldsworthy and
Heathcote (1963) conducted trials in northern Nigeria and found no re-
sponse to K application. Later, Heathcote (1972) found that, under con-
tinuous cropping, groundnut responded to K in the third and fourth years
but not in the first two years. This was consistent with the idea that
groundnut removes relatively small amounts of K and will respond to K
once soil K is reduced to low concentrations. Haggin and Koyumjisky
(1966) recorded significant responses to K in only two of 24 fields in Israel.
Oil and protein percentages as well as yield of tropical groundnut crops
responded positively to 15 kg K/ha in two experiments with initial soil K
levels of 0.28 and 0.40 meq K/lOO g as neutral normal ammonium acetate
(Kayode, 1987). However, yield responded negatively to greater rates of K
application in Kayode's study.
Hartzog and Adams (1973a) conducted 34 on-farm K fertilization trials
in Alabama without yield response and concluded that direct fertilization
of groundnuts was not a good practice, but that K fertilization should be
made to the crops grown in rotation with groundnuts. In Ca and K
experiments on sandy soils in the coastal plain of Georgia, Walker et al.
Potassium (K) 231
(1979) found a decrease in yield with increased K fertilization rates of 0,
112 and 224 kg/ha, which resulted in Mehlich 1 soil test K levels of 20, 25
and 91 mg Klha, respectively. On long-term fertility plots, Cope et al.
(1984) found modest but significant response to 19 kg/ha of fertilizer K
over a period of 30 years. Mehlich 1 soil K test remained fairly constant at
44 mg/kg during the 30-year period. They concluded that the relative
response of multiple crops to soil (and fertilizer) K levels was: cotton>
grain sorghum> corn> soybean> wheat and groundnuts.
Potassium levels in the soil and applied K should be considered in
relation to levels of other cations, especially Ca, as they compete for
uptake by the developing pods. This subject is covered in section
7.2.1.
Indications of optimal soil K levels for yield and quality vary according
to the extractant used, which in turn is determined by region and soil type.
For the Mehlich 1 extractant used in the runner-type peanut belt of the
southern coastal plain of the USA, the identified critical level is 11-13 mg
Klkg (Walker et al., 1979; Mitchell and Adams, 1993). The minimum
sufficiency level used in K recommendation is 20 mg Klkg in order to
provide 'insurance' of adequate available K, bearing in mind sampling and
analytical errors (Mitchell and Adams, 1993). Cox (unpublished studies)
evaluated the Mehlich 3 extractant for use in the virginia-type peanut belt
of North Carolina and determined a critical level between 39 and 47 mg
Kll. However, correlations of yield response to surface soil K levels are not
high in the soils of the peanut belt of the south-east USA because much of
the K not utilized by crops accumulates ill the subsoil. Some of this K is
accessible to the deep roots of the groundnut. Therefore a low soil test
from the top soil does not take into account the K available in the subsoil,
and can underestimate K availability and recommend K fertilization when
there is scant chance of response. Recognition of the unresponsiveness of
the groundnut has recently directed the states in the south-eastern runner-
type peanut belt of the USA to reduce application of K by recognizing a
lower critical level in the Mehlich 1 soil test K of 20 mg/kg (Mitchell and
Adams, 1993).
Leaf K has not been used to any large extent as a guide to fertilization of
the groundnut since the timing of such a diagnostic tool is generally too late
for making any correction. Also, as reviewed above, the problem may be
high rather than low leaf K, thus leaving little that may be done to make a
correction. The exception is to add a soluble Ca source, such as gypsum,
when the KlCa ratio is high. However, leaf K concentration can serve as a
diagnostic for future groundnut fertilization in fields with a similar history.
Since the review of Cox et al. (1982), Walker et al. (1989) have determined
that, for maximum yield of runner-type groundnuts, the minimum suffi-
ciency levels in youngest mature leaflet samples at approximately 100 days
following planting should be 1.0% K. However, normal diagnostic tissue
sampling should occur much earlier (Table 7.2).
7.6 MAGNESIUM (Mg)
Little response has been recorded for application of Mg to groundnuts.

Cox et al. (1982) stated: The literature is almost barren of reports that
peanut yields increased by Mg fertilization.' Responses only occur at very
low soil test levels and they are most likely on deep, excessively drained
sands. Since 1982, Hartzog and Adams (1988b) have reported no response
to MgS04 on a McLaurin loamy sand with a Mehlich 1 Mg test of
3.5 mg/kg. Walker et al. (1989) found no response to Mg on a Fuquay sand
with an initial Mehlich 1 Mg test of 7 mg/kg, but application of 67 kg Mg/ha
increased yield by an average of 15% over the three years of the study on a
Lakeland sand with an initial soil test of 4 mg Mg/kg. In the latter study,
both K and Mg were deficient and application of both increased yield by
69% over the no K and no Mg control. It is very likely that the deep
rooting pattern of the groundnut allows the plant to forage deeply for Mg
as well as for other nutrients. As mentioned for K, some soils retain greater
concentrations of Mg in the subsoil than in the surface horizon. Such is the
case for the Fuquay sand while the Lakeland soil has essentially no
colloidal matter in the rooting zone. The findings of Walker et al. (1989)
therefore seem consistent with the subsoil retention explanation for the
lack of response to fertilizer Mg in soils that have had dolomitic lime
applied in the past, but have low soil test Mg just prior to groundnut.
Schmidt and Cox (1992) did not find a yield response that could be
attributed to Mg fertilization in North Carolina soils with soil tests as low as
0.02 cmolll (approximately equivalent to 5 mg/kg). Their soil tests were
with Mehlich 3, which removes about the same concentration of Mg as
Mehlich 1 (Gascho et al., 1990; Hanlon and Johnson, 1984).
Soil analysis may be useful for prediction of Mg needs if the whole of the
rooting depth is sampled. However, such samples seem impractical and
surface samples are probably not reliable. Walker et al. (1989) suggested a
Mehlich 1 extractable level of 11 mg/kg in the topsoil. For the present, the
sufficiency level is set at 15 mg/kg in the runner peanut belt of Alabama,
Florida and Georgia (Hodges et al., 1993). Schmidt and Cox (1992)
indicated that sufficient leaf Mg was attained when Mehlich 3 Mg in the soil
was as low as 0.06 cmolll (approximately equivalent to 15 mg/kg). At this
low concentration, Mg was only 3% of the cation exchange capacity of the
soil. The actual critical level has not been attained in most studies and is
often complicated by the groundnut plant's ability to forage deeply in soil
horizons which may supplement the surface horizon in supplying Mg.
Use of leaf Mg concentrations for the groundnut suffers from all the
drawbacks listed for K above, including variation with plant age. Walker et
al. (1989) suggest a critical concentration of 0.2% in the recently mature
leaflets of runner-type varieties sampled at approximately 100 days after
planting. Schmidt and Cox (1992) found no Mg deficiency with leaf concen-
trations as low as 0.15%. As for soil analysis, the lower limits of the
Sulphur (S) 233
sufficiency range have not been reached in enough studies to provide a firm
critical level (Table 7.2).
Magnesium may also interfere with Ca uptake by the developing pod;
however, published evidence is scanty. Gascho et at. (1992) found that
germination percentage of seed is somewhat better related to a ratio of
Ca/(Ca+K+Mg) in the nut than to Ca concentration.
7.7 SULPHUR (S)
Even though sulphur is a required nutrient for groundnut, reports of

responses to S applications are scarce in recent literature and are often
confounded by the fact that elemental S application not only supplies S but
also decreases soil pH, thereby changing the availability of other nutrients.
Also, application of S as gypsum supplies Ca, an element of paramount
importance in the groundnut (section 7.2). Recent reports from India
indicate some yield responses on calcareous soils to S applications as
ammonium sulphate, single superphosphate, elemental S and gypsum
(Maliwal and Tank, 1988; Bahl et at., 1986; Sahu and Singh, 1987; Hago
and Salama, 1987). Bahl et at. (1986) found that application of both Sand
Zn resulted in a synergistic effect on increased yield while the main effect
in the studies of Hago and Salama (1987) was to increase chlorophyll a and
b in an area where groundnuts were chlorotic due to Fe deficiency. These
results are attributed to decreased soil pH due to S application. Cox et at.
(1982) provided a review of reports of S responses in groundnut growing
areas of the world and discussed the complications of the reports due to
interactions with pH, P and micronutrients.
In the past, S was supplied to most groundnuts incidentally due to the
application of S-containing fertilizers such as single superphosphate. How-
ever, most modern fertilizers, such as triple superphosphate, are concen-
trated in the major nutrients and contain little S; therefore, more S
deficiency is expected to appear with time in many crops. Groundnuts are
probably unique, as much of the crop receives gypsum to supply Ca and
thus adequate S is provided incidentally. In addition, several of the fungi-
cides used in the past contained S, but use of these for groundnuts has now
decreased.
Atmospheric S is deposited on soils from the burning of fossil fuels. Such
deposits are particularly significant in industrialized areas and are usually
sufficient for crop needs. Common levels of S deposits from the atmos-
phere range from 10 to 20 kg/ha/year. Sulphur concentrations can also be
significant in irrigation waters, thus eliminating possibility of any response
to application (Cox et at. 1982).
Sulphur deficiency is most likely on very sandy soils which possess little
anion exchange capacity and will not hold the sulphate anion. The depth of
the sand is an important factor, since clay accumulation in deeper horizons
within the rooting zone will tend to hold the sulphate anion and will lessen
the possibility of S deficiency.
Soil tests for S are not important in groundnuts as critical levels have not
been established. Such an effort would probably not be rewarding: the soils
with S deficiency problems are those where S is leachable, and so S will be
too elusive for any diagnostic soil test.
Several attempts have been made to establish critical levels of S in the
groundnut plant. Although the literature is not consistent with regard to a
specific S critical level, in general the S concentration should be similar to
the P concentration. The S concentration is also related to the N concen-
tration and an N:S ratio of about 15:1 is desirable, based on a balance of
S-containing and non-S-containing amino acids (Bockelee-Morvan and
Martin, 1966; Lund and Murdock, 1978).
7.8 MICRONUTRIENTS
Micronutrient availability in soils for the groundnut and other plants is

related to soil pH as well as soil physical and other chemical characteristics.
Most micronutrients are more likely to be deficient at high pH, particularly
in calcareous soils. Micronutrients in this category are manganese, zinc,
iron and copper. Manganese and zinc may occur at toxicity levels in very
acid conditions. In contrast, molybdenum is less available in acidic soils.
The single most important consideration in proper availability of micro-
nutrients for plants is soil pH. Secondly, cation and anion exchange
capacity can affect leaching and critical toxic levels in the soil. Thirdly,
interactions of nutrients can result in nutrient imbalance (e.g. a low Ca:Zn
ratio may induce Zn toxicity). Fourthly, the ambient concentrations of
micronutrients in soils and in irrigation waters vary due to soil formation
factors and to previous cropping history.
7.8.1 Boron (B)

Boron can be important when present at either deficient or toxic levels.
The major problem in groundnut is deficiency on highly weathered sandy
soils due to their inability to retain the mobile-borate anion. All groundnut
growing states of the USA recommend boron application to the crop.
Perry (1971) recommended 0.6 kg Bfha for sandy soils and 1.1 kgfha for
heavy soils, but warned against over application in view of potential B
toxicity.
(a) Deficiency
A deficiency most often results in internal nut damage termed 'hollow
heart' which greatly reduces the quality and value of the crop. The con-
Micronutrients 235
dition refers to cotyledons which are small and discoloured. Early research
in Florida found that B deficiency resulted in hollow heart, compacted
branch terminals and cracks on pods (Harris and Gilman, 1957). These
symptoms were first described in greenhouse studies and were later evalu-
ated in field studies with B application in several growing areas.
Application of 1.9 kg H3B03/ha increased yield and grade in the green-
house, but no B deficiency was detected in field studies. Harris and Gilman
(1957) also noted differences in B response between runner varieties.
Harris (1963) described B deficiency in cv. Florigiant groundnuts as hollow
heart, with a mosaic pattern on the foliage. Boron deficiency symptoms are
similar for runner, spanish and virginia types (Harris, 1965). Harris and
Brolmann (1966a) showed that B deficiency symptoms were accentuated
by application of complete fertilizer without B. Deficiency affects the
flowering pattern by producing a less intense but longer flowering period
(Harris and Brolmann, 1966b). Harris and Brolmann (1966c) illustrated
the influence of B deficiency on the cotyledons and showed that plumules
were discoloured and plumule tips were pointed or poorly developed.
Harris (1968) indicated that B should be applied at soil B ~0.2 mg/kg. He
also noted that spanish and runner types developed B deficiency symptoms
earlier than the virginia type, and that the virginia type recovered more
quickly after B application. Shiralipour et al. (1969) determined that B
deficiency resulted in increased nitrogen and amino acid contents in
groundnut leaves without affecting protein levels.
Research in North Carolina using sand cultures showed that B deficiency
resulted in deep green, mottled leaves and terminal death with extensive
secondary branching and decreased internode length (Reid and York,
1958). Cox and Reid (1964) found that 0.6 kg B/ha decreased plumule
damage in a field study. They also showed that liming increased soil
available B, but did not increase B content in groundnut kernels. Stoller
(1966) illustrated the influence of long day length and high light intensity
on accentuating B deficiency symptoms.
In Virginia, hollow-heart symptoms were noted in 1958 but were not
confirmed to be B deficiency until 1965 (Anon, 1965). Research showed
that 1.1 kg B/ha decreased damage, but 2.2 kglha could be toxic. Hallock
(1966) also showed marked decrease in hollow heart by B application, but
rates of 1-2 kg B/ha could be phytotoxic. He also stated that B deficiency is
more common in sandy, droughty soils. Allison (1980) recommended
0.6 kg B/ha application at early bloom.
In Alabama, Hartzog and Adams (1968) determined that topdressing
1.1 kg B/ha had no effect on yield, and increased grade in only one out of
five experiments. Hartzog and Adams (1971) reported that, in eight experi-
ments with hot-water extractable soil B <0.07 mglkg, hollow heart failed
to develop, and yield and grade were unaffected by B fertilization. Hartzog
and Adams (1973a) again reported no yield or grade effect of B fertiliz-
ation, and determined that B could be applied preplant in a herbicide tank
mixture or sprayed on with fungicide without damaging plants. Cope et al.
(1984) reported that yields were not affected by application of B (in
combination with Zn, Mn, Cu and Mo).
Snyman (1972) in South Africa and Morrill et al. (1977) in Oklahoma
determined that application of fertilizer B nearly eliminated hollow heart.
In some cases outside of the USA, deficiency has also been reported to
reduce yield. In India, Saxena and Mehrotra (1985) found a significant
response in yield to the application of 1 kg B/ha on a loam soil in a trial
where up to 2 kg/ha was applied. In similar trials on a sandy loam they
found yield response to only 0.5 kg B/ha. Yields in an experiment in China
(Zhang et al., 1986) were increased by up to 11.5 % by application of B (as
borax). On the other hand, Blarney et al. (1981) in South Africa empha-
sized the narrow window between deficiency and toxicity. They found
decreased growth and yield by residual and applied B applications on a
sandy loam soil with an initial pH of 3.9 (0.1 N KCl). Annual B appli-
cations of 1 kg/ha significantly reduced both kernel and hay yields.
Applications of limestone at rates of up to 2.4 tons/ha in previous crops did
not reduce toxicity problems, suggesting that the increased root growth
with higher pH increased the uptake of B0 3 enough to offset any fixation
of B due to precipitation of Al(OHh
In India, Rao et al. (1960) reported that 11 kg B/ha did not affect yield.
Gopal and Rao (1972) also found no benefit from B application but gave a
critical level of 25 mg/kg in middle leaves. Other studies showed that: 9 kg
B/ha increased yield and oil content (Ganesan and Sundararajan, 1972);
15 kg/ha borax increased yield (Muthuswamy and Sundararajan, 1973);
6 kg/ha borax or boric acid increased yield and quality (Asokan and Raj,
1974); B application increased oil content (Sankaran et al., 1977); B spray
had no yield effect (Swamy and Reddy, 1983); 2 mg/kg B gave maximum
yield (Golakiya and Patel, 1986); pod yield was highest when Ca:B ratio in
pods was 218-224 (Golakiya, 1989); and 5 kg B/ha increased yield on a soil
with 0.16 mg B/kg (Jadhao et al., 1989). Boron deficiency was also reported
in Malawi on light, sandy soils (Anon, 1972).
Research on spanish-type groundnuts in Oklahoma suggested a critical
level of 30 mg B/kg in young leaves (Chrudimsky, 1970). Hill and Morrill
(1974) found B deficiency in 50% of their field locations but reported that
B application did not affect yield or grade. They stated that hollow heart
was related to soil B (hot-water soluble) <0.15 mg/kg and leaf B of
26-30 mg/kg. Hill and Morrill (1975) found that B application improved
kernel grade, except at high potassium rates. Morrill et al. (1977) con-
cluded that soil B ~0.15 mg/kg and leaf B <30 mg/kg require B fertiliz-
ation at a rate of 0.6 kg B/ha.
Diagnosis of B nutritional problems is by both soil and plant analysis but
neither has been entirely satisfactory. Reliability of soil B tests has been
considered rather low, due mostly to the fact that few correlation data have
been produced in recent years. The critical values (above which there
Micronutrients 237
should be no deficiency) of 0.05 mg B/kg established by Cox and Reid

(1964) for acid soils and 0.2 mg B/kg by Hill and Morrill (1974) for
calcareous soils by the hot-water soluble method are the best estimates
available. In the south-eastern USA 0.6 kg B/ha is recommended when soil
B <0.2 mg/kg.
The critical leaf B concentration, below which spanish-type ground nut is
deficient, is in the range 25-30 mg/kg (Morrill et al., 1977). Toxic concen-
trations reported for spanish and virginia types vary from 60 to 112 mg/kg
(Gopal, 1971a, 1971b; Luke, 1969; Blarney and Chapman, 1979). In the
south-eastern USA, 0.6 kg B/ha is recommended if leaf B is <30 mg/kg
(young leaves). Very little B-related research has been reported in the past
decade. There is a need for studies to evaluate diagnostic B concentrations
using modern testing methods such as inductively coupled plasma (ICP)
and/or direct current plasma (DCP) emission spectroscopy methods, which
are less laborious.
Boron deficient conditions in sandy soils can be alleviated by foliar
applications. Walker (1967) stated that 0.6 kg B/ha applied as a foliar spray
increased yields on Ruston and Tifton soils but not on Greenville soil (a
finer textured soil). Application of a total of 0.6 kg B/ha is commonly tank-
mixed with and split equally between the first two fungicide applications.
(b) Toxicity
Boron can be toxic to groundnuts; therefore, it should be applied at the
recommended rate only. McGill and Bergeaux (1966) warned of exceeding
0.6 kg B/ha, and Stoller (1966) showed that the critical toxicity level for B
was 100 mg/kg in shoots. Morrill et al. (1977) stated that 1-1.5 kg B/ha
caused toxicity and reduced yields. Application of 1 kg B/ha reduced yields
in Australia (Blarney et al., 1981). A 10% yield decline was associated with
58 mg B/kg in young leaves. In Israel, B concentration of 0.29 mM in
nutrient solution decreased yield (Lauter et al., 1989).
Research in India has shown that B toxicity results in chlorosis of leaf
tips which extends marginally and interveinally, followed by marginal
necrosis (Harigopal and Rao, 1964). Chlorosis was related to decreased
leaf chlorophyll, protein N, and Fe. Boron toxicity decreased yield, and
leaching was recommended as an ameliorative q1ethod (Harigopal and
Rao, 1967). Gopal (1968) stated that 10 mg B/I in solution and 85 mg/kg in
leaves were related to toxicity. Gopal (1969) showed that B interfered with
the ability of Fe to complex with proteins. However, Fe addition in a pot
study made no difference (Gopal, 1970a). Boron content of middle leaves
of toxic plants was> 1100 mg/kg. Boron also decreased leaf Cu (Gopal,
1970b). Gopal (1971a) found 3 mg B/kg in soil to be toxic, and Gopal and
Rao (1972) found 140 mg/kg to be the critical level in middle leaves. Boron
toxicity decreased total N and protein N in chlorotic leaves, but increased
soluble N and free amino acids (Gopal, 1971b). Boron application at
>6 kg/ha borax gave an adverse effect (Asokan and Raj, 1974), and
5 kg/ha borax resulted in toxic symptoms (Reddy and Patil, 1980). Care
should be taken not to overapply B to groundnuts.
7.8.2 Chlorine (CI)

Chlorine toxicity has been described for soybeans in Georgia (Parker et al.,
1983) but has not been found in groundnuts. Chlorine is an essential
element for plant production but CI deficiency has not been described for
groundnuts.
Schilling and Hirsch (1974) found no correlation between leaf CI and
groundnut yield in a study in Senegal. Chloride effects on cv. Florunner
were studied in the greenhouse and field in Georgia (Parker et al., 1984).
Addition of CI to soil increased CI concentration in leaves but there was no
significant effect on dry matter production (greenhouse) or pod yield
(field).
There are no data that would warrant fertilizer CI recommendations for
groundnuts.
7.8.3 Copper (Cn)

Copper is sometimes applied to agricultural crops as a micronutrient but is
commonly applied in the form of pesticides, particularly fungicides.
Bledsoe and Harris (1947, 1948, 1949) reported that application of 11 kg
CuCl2/ha increased the proportion of sound to shrivelled nuts for runner
groundnuts. Three years after application, the residual effect of Cu on
quality was maintained. Harris (1952) described Cu deficiency symptoms
as affecting the bud area in particular, as well as causing small, irregular
leaflets with marginal necrosis and mild chlorosis and small yellow-white
spots on the foliage. Harris (1952) found that the spanish type is more
sensitive to Cu deficiency (han the runner type, but that yields for all three
cultivars studied (two runners and one spanish) were increased more than
300% by applying 11 kg/ha Cu (as CuC}z) to an Arredondo loamy fine sand
(pH 5.7). Copper application also decreased seed shrivelling and increased
the percentage of sound mature kernels (SMKs). The residual effect of soil
Cu application (11 kg/ha) to oats, wheat, rye or cotton in rotation with
groundnuts was found to be equally effective as groundnut foliar appli-
cations (0.2 kg CuCI2/ha). However, Harris (1952) concluded that, in
general, yields in Florida had not been increased by Cu applications
(though yields were increased on the Gainesville experimental farm) and,
therefore, Cu application was not recommended.
In research in Georgia, Boswell (1964) stated that no definite pattern
was found between Cu application and yields. Gopal (1970a) reported that
Micronutrients 239
B treatment decreased leaf Cu concentration and Cu-protein enzyme
activity in groundnuts. Gopal (1975) stated that 10 mg/l B in sand culture
decreased leaf Cu concentration by 24%. Katawatin et al. (1989) deter-
mined that phosphorus fertilization decreased Cu uptake.
7.8.4 Iron (Fe)

Iron deficiency can be a serious problem in calcareous soils but no prob-
lems have been reported on acidic soils. Perkins (1964) stated that the total
Fe content of most acidic soils in Georgia is greater than 1%; therefore he
assumed that Fe is available in Georgia in sufficient amounts for crop
production. Iron deficiency results in interveinal chlorosis (starting in the
youngest leaves), followed by chlorosis of the entire leaf (whitish-yellow)
and brown spots leading to marginal necrosis. Lachover and Ebercon
(1972b) showed that yield response to Fe application is related to
CaC0 3 concentration in the soil. Papastylianou (1989) surveyed 35
groundnut fields in Cyprus and determined that plants were chlorotic in
soils with CaC0 3 >20-25% and when Fe content was less than 2.5 kg/ha.
Dungarwal et al. (1974) applied 500 kg S/ha to a clay loam of pH 8.4 and
increased yields by 197% evidently due to reduced pH and increased Fe
availability.
Lachover et al. (1970) applied an iron chelate (FeEDDHA) to a soil in
Israel with pH 7.9 and 15% CaC0 3 and measured a 50% increase in pod
yield and a 40% increase in hay yield. Lachover and Ebercon (1971)
showed that Fe chelate applied to a soil of pH 7.9 and 11 % CaC0 3 caused
leaves to green up and increased yield. Foliar applied Fe chelate increased
greenness and yield, but Fe polyflacenoid and Fe acetate increased green-
ness without improving yield (Hartzook et al., 1971). Yields were increased
359% by application of 10 kg Fe/ha (as FeEDDHA) to a loamy clay with
pH 7.9 and 31% CaC0 3 (Lachover and Ebercon, 1972a).
Reddy and Patil (1980) applied FeS04 spray to spanish-type groundnuts
grown on an Indian soil with pH 7.5 (2.5% CaC0 3 and 9 mg/kg Fe) and
measured no yield increase. Hallock (1964) applied Fe chelates to ground-
nuts grown in Virginia and found no yield effect. Schneider and Anderson
(1972) measured yield response to FeEDDHA in Texas.
Patil et al. (1979) determined that foliar application of FeS04 produced
greater yields than soil applied FeS04 on a black clay soil with pH 7.7
(2.5% CaC0 3 and 1.26 mg/kg available Fe). They also noted that high
phosphorus fertilization was related to chlorosis and diminished yields.
Nitrogen fertilizers did not improve leaf colour in groundnuts with Fe
chlorosis and N levels in green and chlorotic leaves were not different
(Lachover and Ebercon, 1971). Kafkafi and Neumann (1985) showed that
no Fe chlorosis was observed when NH4-N was 20% or greater of the total
soil N. They suggested that NH4 + uptake resulted in H+ efflux from roots
which reduced the soil Fe 3 + to Fe 2 + which groundnut plants then absorbed.
Kafkafi and Neumann (1985) recommended the use of a nitrification
inhibitor in addition to ammonium fertilizers. Iron stress was found to be
stronger in nodulated than in non-nodulated plants and high plant Fe levels
enhanced N fixation (Terry et al., 1988).
Zaharieva et al. (1988) showed that high soil Mn levels can be related to
Fe deficiency. Iron and Mn do not compete for absorption sites; Mn
actually inactivates Fe metabolic activity by decreasing the Fe2 + concen-
tration in groundnut plants. Zaharieva et al. (1988) also stated that Fe
efficient cultivars can overcome Fe chlorosis without additional Fe supply.
Hartzook et al. (1974a,b) determined that yield of untreated Fe efficient
cultivars was roughly equal to the yield of inefficient cultivars which were
fertilized with an Fe chelate.
Iron deficiency cannot be determined by plant analysis since chlorotic
leaves may contain more Fe than healthy leaves (Papastylianou, 1989).
Rao et al. (1987) stated that total Fe is unsatisfactory as a measure of Fe
status in plant tissue, and that Fe status was better assessed from an
estimate of ferrous-Fe (by extraction with o-phenanthroline). They deter-
mined that the critical level for ferrous-Fe in the youngest fully opened leaf
was 6 mg/kg. Rao et al. (1987) showed that chlorotic leaves had lower
extractable Fe but higher total Fe, so that Fe deficiency is due to poor
utilization of Fe within a leaf, not absorption or translocation problems.
Estimated critical levels are 2.5 mg Fe/kg in soil and 6 mg/kg ferrous-Fe
in leaves.
7.8.5 Manganese (Mn)
( a) Deficiency
Manganese deficiency is a problem only on high pH soils. Rich (1956)
stated Mn deficiency is a problem for the crop in Virginia. He reported that
Mn concentration in the plant was inversely related to soil pH, calcium,
and magnesium levels, in a study using 32 Coastal Plain soils. Visual Mn
deficiency symptoms were evident when leaf Mn was below 10 mg/kg
(Rich, 1956). Manganese deficiency in groundnut has been observed on
soils with pH values as low as 5.8. Anderson (1964) reported that research
in Georgia showed no yield effect of MnS04 additions (10-50 kg/ha) to a
Tifton loamy sand with pH 6.5, a Norfolk sandy loam or a Greenville clay
loam. Hickey et al. (1974) recorded significant yield increase for crops
grown on a Lakeland sand (pH 6.3, extractable soil Mn 0.67 mg/kg) due to
addition of 40 kg Mn/ha (MnCh). Foliage Mn levels were also increased
from 59 to 155 mg/kg. The recommendation from the 1980 Virginia Peanut
Production Guide is to apply foliar Mn at a rate of 0.8-1.1 kg/ha in each of
up to three applications, when interveinal chlorosis, symptomatic of Mn
deficiency, is evident (Allison, 1980).
Micronutrients 241
Hallock (1979) showed that soil Mn treatments did not yield significantly
less than spray treatments for virginia-type groundnuts but deficiency
symptoms were greater for soil applied treatments. Parker and Walker
(1986) studied the interaction of Mn response with soil pH on a Pelham
sand. Manganese deficiency occurred on plots with pH levels near 6.8
(Mehlich 1 soil Mn = 3.3 mg/kg) but not in plots with pH levels of 5.2 (soil
Mn = 2.1 mg/kg) or 6.0 (soil Mn = 2.5 mg/kg). At pH 6.8, soil application
of Mn at 0, 10, 20 and 40 kg/ha resulted in yields of 3.41, 5.40, 5.73 and
6.37 tonnes/ha, respectively. Critical Mn levels in the leaves were 13, 15,
15,15,13 and 12mg/kg at 7, 9, 11,13,15 and 17 weeks after planting.
Parker and Walker (1986) concluded that maintaining a soil pH near 6.0
was optimal for groundnut production.
Soil Mn applications can be used to prevent Mn deficiency when the soil
pH is known to be >6.0. Foliar Mn applications can correct Mn deficiency,
diagnosed through foliar symptoms, more rapidly than soil Mn applica-
tions.
(b) Toxicity
Manganese toxicity can be a problem in low pH soils. Morris and Pierre
(1949) reported that a concentration of 10 mg Mn/l in a nutrient solution
reduced growth to 76% of the control and resulted in chlorosis of leaf
margins. Plant Mn concentration was 1245 mg/kg. Groundnuts were the
least sensitive (of five legumes studied) to Mn toxicity and had the lowest
plant Mn concentrations. Boyd (1971) described Mn toxicity symptoms for
the crop as interveinal leaf chlorosis followed by marginal leaf necrosis.
Boyd also found that soil Mn (NH 4 0Ac extractable) was well correlated
with leaf necrosis. Severe symptoms occurred when soil Mn was greater
than 10 mg/kg and foliar Mn was greater than 50 mg/kg.
Benac (1976) found that a nutrient solution with Mn concentration
~20 mg/kg caused stunting of plants. Tissue Mn of plants with leaf necrosis
was ~4000 mg/kg, and the highest Mn concentration was found in the
leaves. In a nutrient solution test Nambiar and Anjaiah (1989) found that
groundnut plants with Mn toxicity symptoms had between 1040 and
3070 mg/kg Mn in the plant tissue. High Mn levels decreased dry matter
accumulation and nitrogen uptake.
High Mn levels may magnify Fe deficiency. Zaharieva (1986) found
that the Mn:Fe 2 + ratio in plants was greater than 1:1 in chlorotic
plants. Zaharieva et al. (1988) stated that groundnut plants with Mn
toxicity (dark brown marginal leaf spotting) had >450 mg/kg Mn in the
leaves. Application of Fe as FeEDDHA eliminated leaf spotting.
Zaharieva et al. (1988) also suggested that Fe application could induce Mn
deficiency.
It is difficult to relate nutrient solution tests to field situations. More
research is needed in the area of Mn toxicity in groundnuts.
7.8.6 Molybdenum (Mo)
Molybdenum is essential for nitrogen fixation and is therefore recom-
mended for some legumes (e.g. soybeans, alfalfa). However, it is currently
not recommended for groundnuts.
Plot trials in Senegal showed that Mo had a harmful effect on groundnuts
(Bouyer and Collot, 1952). Molybdenum deficiency symptoms were not
obtained in a sand culture experiment, though slight chlorosis was evident
late in the growing season (Reid and York, 1958). Harris (1959) stated that
Mo application caused foliage to be a darker green and frequently
increased the size of the foliage, but it has never caused a significant
increase in yield in research in Florida.
Rao et al. (1960) reported that an application of 140 g Mo/ha in India
increased pod yield. Walker (1967) found that 0.2 kg Mo/ha increased yield
by 224 kg/ha on a Tifton soil but had no effect on yield on a Greenville soil.
Welch and Anderson (1962) found that Mo availability was increased by
liming and that Mo application increased Mo concentration in leaves, but
no deficiency symptoms were evident in areas which received no Mo.
They stated that seed Mo concentration is high enough to provide the
plant's Mo requirement even in a low Mo soil. Sellschop (1967) stated that
Mo deficiency is best corrected by liming. Parker (1964) reported that
Mo often improved plant colour but gave a yield response in only one of
15 experiments conducted in Georgia. Boswell et al. (1967) showed that
yield was not well correlated with leaf or soil Mo content, and Mo addition
increased nitrogen content of foliage. However, the yield effect of Mo was
inconsistent.
Heinis (1972) determined that Mo fertilization increased N and methion-
ine contents in leaves. Graham (1979) stated that Mo is essential for nodule
formation and function and Mo deficiency, which is more common on acid
soils, can cause nitrogen deficiency. Kiat (1979) reported that Mo appli-
cation had no significant influence on yield, nitrogen fixation or Mo
concentration in groundnut tissue.
In research in India, it was found that 1 kg/ha NH4 molybdate increased
yield of spanish groundnut (Reddy and Patil, 1980). The soil test level was
0.5 mg/kg available Mo, and pH was 7.5. The authors suggested that this
beneficial effect may be due to increased N availability which resulted in
increased protein in groundnut kernels. Kene et al. (1988) found that Mo
increased nodulation and nodule N content for groundnut.
Most of the literature agrees that Mo increases greenness and N content
of groundnut leaves, but yield increases due to Mo application are rare.
More research is required to determine under what conditions Mo ferti-
lizers may be beneficial. Currently, there is no consistent data to recom-
mend Mo fertilization for groundnuts.
Micronutrients 243
Zinc (Zn)
(a) Deficiency
Zinc deficiency usually occurs under high pH conditions. Carter (1964)
summarized Georgia research and showed that, even though zinc fertiliz-
ation sometimes increased yield and sometimes decreased yield, in general
the differences were not significant. Sellschop (1967) stated that Zn insuffi-
ciency was less conspicuous in groundnut than in maize, and recommended
16-22 kg Zn/ha where the problem is common. Schneider and Anderson
(1972) found that a Zn application of 0.1 kg Zn/ha gave a positive yield
response for spanish groundnut, but that 90 kg/ha reduced yield. In a
calcareous soil with <0.3 mg/kg soil Zn, applications of ZnS04 from 10 to
60 kg/ha had no significant yield effect (Lakshminarasimhan et al.,1977).
Spraying 0.8% ZnO on a calcareous vertisol increased yield of valencia-
type groundnut (Mupawose, 1978). Phosphorus application can show an
antagonistic effect on Zn uptake (Chahal and Ahluwalia, 1977). Zinc
deficiency is associated with high soil pH and high available P levels
(Graham, 1979). However, Patil et al. (1979) applied ZnS04 to groundnuts
with severe chlorosis, which was attributed to high soil pH and heavy
phosphorus fertilization, but there was no yield response to either soil or
foliar applications.
Reddy and Patil (1980) stated that 0.5 mg/kg Zn in soil and 22 mg/kg Zn
in leaves at flowering were the critical levels for Zn deficiency in ground-
nuts. Rhoads et al. (1989) applied Zn to soil in a greenhouse study and
determined that the cultivar Southern Runner was more sensitive to Zn
deficiency than Sun runner. They suggested a critical soil Zn (Mehlich 1)
level of 2.5 mg/kg at soil Ca >400 mg/kg.
Bell et at. (1990) described Zn deficiency symptoms in groundnuts as
decreased internode length and restricted development of new leaves.
They also found that Zn deficient plants accumulated reddish pigments in
stems, petioles and leaf veins. They stated that 20 mg/kg in upper stems
and leaves and 25 mg/kg in recently matured leaves (at early pegging) had
been used previously as critical levels for Zn deficiency diagnosis. They
recommended that the blade of the youngest fully expanded leaf be used
for diagnosis with 8-10 mg/kg Zn as its critical value. Zinc deficiency is
related to high soil pH, high soil Ca and high soil P. Leaf critical level is
20-25 mg/kg, and foliar application is probably the best way to correct Zn
deficiency.
(b) Toxicity
Zinc toxicity was first reported by Quintana (1972) who noted that appli-
cation of 90 kg Zn/ha as ZnS04 decreased yields and resulted in 67 mg
Zn/kg in plant tissue. Keisling et al. (1977) described Zn toxicity symptoms
as chlorosis, stunting, purple coloration of the main stem and petioles
usually a lesion at the base of the plant (stem splitting) and premature
necrosis. Tentative Zn toxicity critical values were set at 12 and 220 mg/kg
for soil (Mehlich 1 extractable) and tissue, respectively. Tissue Zn concen-
tration increased 15 mg/kg for each 1 mg/kg increase in soil Zn. Liming
reduced Zn uptake and stunting but did not change the level of Mehlich 1
extractable Zn in soil. Davis-Carter et at. (1990) showed that leaf chlorosis
and stem purpling were not well correlated with leaf Zn levels, and
described new symptoms of horizontal leaf growth and leaf closure.
Rhoads et at. (1989) stated that peanut response to Zn appeared to be
more dependent on soil Ca level than on soil pH. Up to 23 kg Zn/ha did not
affect plant growth at soil Ca >400 mg/kg with soil pH of 6.5 to 6.8, but
8.1 kg Zn/ha reduced plant growth when soil Ca ranged from 150 to
200 mg/kg and pH was ~6.6. Parker et at. (1990) studied data from
growers' fields which indicated that a leaf Ca:Zn ratio ~50 was required for
Zn toxicity to groundnuts. Leaf Zn was affected more by soil pH than by
soil Zn. A regression equation, including both factors, showed that an
increase in soil Zn from 1 to 10 mg/kg increased leaf Zn by 202 mg/kg at
soil pH 4.6 but only 9 mg/kg at pH 6.6. Cox (1990) used data from North
Carolina and Georgia to predict plant Zn concentration from soil pH and
soil Zn. Davis-Carter et at. (1991) stated that, since Mehlich 1 extraction of
Zn from soil is not pH sensitive, it is necessary to include soil pH in any
regressions predicting leaf Zn. They used such equations to calculate the
probabilities for the development of Zn toxicity symptoms as a function of
soil pH and soil Zn. Rhoads et at. (1991a) showed that increasing Zn rates
in greenhouse studies decreased Ca concentration in groundnut tissues. In
two greenhouse tests, the Ca:Zn ratio in tissue proved to be a good
diagnostic tool for predicting dry matter yield; in one test, the critical ratio
was 140:1, and in another it was 78:1.
Rhoads et at. (1989, 1991b) also noted cultivar differences in tolerance to
Zn toxicity. Southern Runner had greater dry matter yield and lower plant
Zn concentration than Sunrunner at the same soil Zn level. Davis-Carter et
at. (1990) illustrated the influence of soil texture on critical levels.
Groundnuts grown on clayey soils required lower soil pH and higher soil Zn
levels to develop toxicity symptoms than those grown on sandy soils. On
clayey soils, leaf Zn >470 mg/kg was related to toxicity. However, on sandy
soils, plants with leaf Zn >350 mg/kg exhibited zinc toxicity symptoms.
7.9 CURRENT EMPHASIS AND RESEARCH NEEDS
Only a few scientists are now actively working to improve the mineral
nutrition of groundnut. Many studies appear to be localized and indicate
responses on a particular soil or in a local situation. Often studies provide
only yield results from applications of various fertilizers applied at several
rates, without providing adequate supporting soil and plant analytical
References 245
results. Such studies can certainly provide local help but are difficult to
extend beyond limited confines. Recent emphasis is placed on efficiency of
production. Supplying inputs, such as fertilizers, at rates that do not exceed
needs is consistent with both top economic returns and environmental
concerns. For example, emphasis in the south-eastern USA peanut belt has
recently been placed on reviewing fertilizer recommendations for ground-
nuts. The review indicated that responses to P and K are rare when
groundnut is in a rotation with other crops to which fertilizer is applied.
Reviews such as this can lead to increased efficiency; they also emphasize
the fallacy of making general applications of all or most nutrients without
consideration of plant needs via research-based recommendations.
Another major consideration for groundnut is quality. Seed quality and
aflatoxins in groundnuts are affected by mineral nutrition. Some studies
have attempted to relate quality to mineral nutrition, but more research
may be quite fruitful.
Increasing interest in and need for better waste utilization throughout
the world makes it imperative to study the effects of application of some
wastes to agricultural lands. The effects of several by-products and waste
products on groundnut are unknown but researchable.
In areas subject to erosion, conservation tillage methods are being
adopted rapidly. Studies are needed to define and solve problems unique
to tillage methods that provide maximum residue coverage and minimum
soil disturbance.
Finally, increased linkage between scientists will lead to major improve-
ments. Several possibilities can be cited, but one of them may be the
improvement of N fixation in groundnut by linking breeders with plant
nutritionists. Links between scientists are also needed to study groundnut
nutrition as part of a total cropping system rather than as a single crop.
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CHAPTER 8
Nitrogen fixation
J. Sprent
8.1 INTRODUCTION
Groundnuts belong to the subfamily Papilionoideae of the family

Leguminosae, using the classification system of Polhill and Raven (1981).
Most but not all of the Papilionoideae form nitrogen fixing nodules with
soil organisms known generally as rhizobia (Faria et at. 1989). General
characteristics of advanced papilionoid nodules are listed in Table 8.l.
Variations on this theme are discussed elsewhere (Sprent, 1989; Sprent and
Sprent, 1990) except for characters specific to Arachis which are con-
sidered in the next section.
Within an active nodule, rhizobia carry out the reduction of nitrogen gas
to ammonia using the enzyme complex nitrogenase:
N2 + 8H+ + 8e- ~ 2NH3 + H2
The reduction of protons to hydrogen gas appears to be an inevitable
concomitant to the reduction of N2 to ammonia. Nitrogenase can also
use a number of other substrates, the best known of which is acetylene,
C2H 2: this reaction forms the basis of the acetylene reduction (AR)
assay for nitrogenase. Reactions are coupled to the hydrolysis of ATP to
ADP + Pi, with probable stoichiometry of 16 ATP for reduction of one
molecule to N2. This, coupled with the need for reducing power at a
redox potential of around -430m V, means that the nitrogenase reac-
tion is energy intensive. Further energy is required to assimilate the
products of fixation into amino acids. However, this is true also for
assimilation of soil ammonium and nitrate (Figure 8.1). Many people
have attempted to compare the costs of nitrogen fixation and nitrate
assimilation in theory and by experiment. Results vary with species and
assumptions made, but it is generally accepted that nitrogen fixation is
more expensive to the plant than nitrate assimilation and that both are
more expensive than ammonium assimilation (Raven, 1988; Vance and
Heichel,1991).
256 Nitrogen fixation
TABLE 8.1 Some general features of nodules of members of well-studied tribes of
the Papilionoideae
Group
Character 2 3
Bacterial Rhizobium Bradyrhizobium Bradyrhizobium

partner (except Phaseolus)
Infection Via root hairs and Via lateral root
process involving infection junctions. Infection
threads threads not formed
Nodule growth Indeterminate Determinate Determinate
Infected region Contains uninfected Uniformly infected
(interstitial) cells
Bacteroid shape Pleomorphic Rod-shaped Spherical or rods
Predominant Amides Ureides Amides
export product
Group 1 is typical of thc Vicieae and Trifolieae and is largely bascd on Vicia, Pisum,
Trifolium and Medicago; Group 2 is typical of many genera of the Phaseoleae and
Desmodieac and is largely based on Glycine, Phaseolus and Vigna; Group 3 is generally
typical of the Aeschynomeneae and some Dalbergieae but details given are based solely on
Arachis: variants are described in Sprent et al. (1989).
In addition to the direct biochemical costs of reducing N2 , legumes have

to make an initial investment in nodule material (Sprent and Thomas,
1984) before they can reap any benefit. The construction costs also include
production of mechanisms for controlling the diffusion of oxygen into
nodules. Because nitrogenase is inactivated by free O 2 and yet the bacteria
need O 2 to produce ATP to drive the reaction, nodules must maintain a
high flux of O 2 at a low concentration from the soil atmosphere to the site
of nitrogen fixation. The exact flux required will vary according to environ-
mental conditions (e.g. temperature, water supply), which will affect not
only nodule metabolism but also the rate at which the plant shoot system
can supply nodules with photosynthate. In the few nodules which have
been studied (e.g. soybean and clover) there appear to be both a fixed and
a variable resistance to oxygen transport located in the nodule cortex
(Witty et at., 1986; Figure 8.6). Once past this barrier, oxygen transport is
facilitated by an extensive intercellular space network and by the presence
of the oxygen carrying pigment, haemoglobin, in the host cells surrounding
the peribacteroid units. The appearance of haemoglobin, which gives
active nodules a distinct pink colour, is closely coupled with the onset of
nitrogenase activity. The latter is coupled to the repression, in the bacteria
(now known as bacteroids), of the ability to assimilate ammonia. Instead
Arachis as a nodulating genus 257
SOIL NITRATE
•
uptake1bY '00" ~
transport to
stem and leaves
reduction to
ammonium ------'.~I
AMMONIUM
ASSIMILATION
VIA
GS/GOGAT
\ reduction to
~--- ammonium
(see figure 8.3)
uptake by roots
t
SOIL AMMONIUM
Figure 8.1 Major pathways of nitrogen assimilation in plants (see Figure 8.3 for
terms OS, OOOAT).
the ammonia (or ammonium) passes to the host cell for assimilation. Very
few rhizobia - and none known to infect groundnut - can be induced both
to fix nitrogen ex planta and to assimilate the products of fixation.
8.2 ARACHIS AS A NODULATING GENUS
Within the Papilionoideae, Arachis is classified III the tribe

Aeschynomeneae. In terms of its nodulation this tribe has a group of
unusual and interesting characters, shared only (as far as we know) with a
few members of the tribe Dalbergiae (Sprent et al., 1989).
Arachis does not normally produce 'typical' root hairs, i.e. outgrowths
from epidermal cells, even though it has the genetic material to do so -
apparently normal root hairs can be induced by salt (Sprent and McInroy,
1984). Multicellular hairs are found near lateral root junctions and,
although these are not involved in the infection process in the way that
normal root hairs on legumes such as clover are, mutants lacking these
hairs do not nodulate (Nambiar et al., 1983). The original observations on
nodulation in Arachis (Allen and Allen, 1940) were followed by various
other studies, of which that of Chandler (1978) is the most comprehensive.
The following description is based on Chandler's work. Infection was only
found where cells at the base of the (usually) multicelluar root hairs at the
root junction were swollen. It was suggested that this swelling may contrib-
ute to separation of cells at the root junction, permitting intercellular
passage of rhizobia. Signs of intracellular penetration of both root hairs
and basal cells were seen but these did not develop further. Intercellular
penetration proceeded into the axillary/subtending root interface and some
adjacent cortical cells (detectable by enlarged nuclei) were penetrated by
rhizobia. No organized infection thread, characteristic of the root hair type
of infection of soybean, clover etc., was seen at any stage. Instead, cell wall
material appeared to be broken down and bacteria spread into the host
cell, although surrounded at all times by an electron-dense matrix and a
host-derived membrane. This rather amorphous structure was termed a
zoogloea. When inside the host cell, the bacteria continued to divide
frequently. The net result was a uniform population of infected cells,
without the frequent uninfected interstitial cells characteristic of most
other nodules studied (Figure 8.2).
When first described, the above process was thought to be unusual.
However, on the basis of absence of interstitial cells and infection threads
and the occurrence of nodules in axils of lateral roots, we believe it to be
common in related legumes. The only other cases where the infection
process has been studied in any detail are Stylosanthes (Chandler et al.,
1982) and Aeschynomene (Arora, 1954). These genera show broadly simi-
lar intercellular passage of rhizobia to that of Arachis but with a few minor
variants such as lack of swollen basal cells in Stylosanthes (Sprent et al.,
1989).
All species of Arachis which have been examined form nodules and so it
may be safe to assume that this nodulation is a generic attribute. Within
genera this is usually the case, although there are exceptions, such as in the
mimosoid genera Acacia and Parkia. As with a number of other genera,
non-nodulating mutant forms of A. hypogaea have been generated and
these are useful experimental tools (Giller et al., 1987).
Distribution of nodules on plants of Arachis may vary with subspecies
and variety. A. hypogaea var. hypogaea has a natural tendency to form
Arachis as a nodulating genus 259
Figure 8.2 Photomicrographs of groundnut root nodules (b, bacteroids; ic,

infected cells; vb, vascular bundle): (a) part of nodulated root (note that nodules
are located in the axils of lateral roots); (b) light micrograph of cortex and part of
infected region of a groundnut nodule; (c) electron micrograph of an infected cell.
Cells in the outer layers of the cortex in (b) have larger cells with more intensely
stained walls than those in the inner layers - compare also with Figures 8.5 and 8.7.
Crystals (arrows) in outer layers are similar to the calcium oxalate crystals observed
by Sutherland and Sprent (1984) in certain ureide-exporting legumes. Most of the
infected cells in (b) are vacuolate. The bacteroids in (c) are spherical; the small
black structures (arrows) are probably oleosomes (Jayaram and Bal, 1991). «b)
and (c) courtesy of K. Wilson and K.E. Giller.)
lateral roots on the hypocotyl and 13-18% of its nodules may be located in
this region (Nambiar et al., 1982). Nodules have also been reported by
these workers to be associated with lateral roots arising at the base of the
stem, a property which needs further investigation but is consistent with
the position in the related genus Aeschynomene (e.g. Arora, 1954).
Nodules on roots are arranged in rows according to protoxylem points.
There are major genotypic differences in the vascular anatomy of the
root/hypocotyllstem regions, where the central root stele divides into
discrete bundles (O'Brien and Sprent, unpublished). These differences
may relate to the occurrence of nodules on the root/stem interface.
8.3 THE RHIZOBIAL COMPONENT
The taxonomy of rhizobia is currently undergoing major changes. All early

classifications were based on host plant affinities but, increasingly, molecu-
lar characters such as 16s-rRNA are being taken into account (Young,
1992). These are showing up interesting affinities between rhizobia and
other soil bacteria. Rhizobia from tropical soils have been less intensively
studied than those from temperate soils, making it unwise to generalize too
widely. Growth rate in culture is still a convenient preliminary. The old
categories of 'slow-growing' and 'fast-growing' forms are still useful, but
we now have additional 'extra slow-growing' forms (e.g. Barnet, 1988). Of
those that have been properly categorized, slow-growing forms usually
conform to the genus Bradyrhizobium and fast-growing forms to
Rhizobium. Arachis is usually nodulated by slow-growing forms of the
Bradyrhizobium sp. (cowpea) type, which generally have a wide host range
and occur abundantly (more than 10 000 per gram) in most tropical soils.
Some strains of fast-growing rhizobia can nodulate Arachis with various
degrees of effectiveness (Rupela et al., 1991), although most are ineffective
(Wong et al., 1988). Other legumes may also be nodulated by either fast- or
slow-growing rhizobia and nodule development is different in some, such
as Lotus (Wood et al., 1985). Ineffective nodules formed by Rhizobium
NGR 234 were found by Wong et al. (1988) to develop in a manner similar
to effective nodules formed by Bradyrhizobium apart from bacteroid
shape, which was polymorphic in the former and spherical (the norm for
ground nut nodules) in the latter.
The widespread occurrence and promiscuity of rhizobia nodulating
Arachis makes difficulties for field studies and inoculation programmes.
Fortunately strains can sometimes be distinguished serologically and mor-
phologically, enabling studies on competition and hostlrhizobial inter-
actions. One such study was carried out by Hadad and Loynachen (1986).
Colony morphology was used as a basic criterion for separating strains,
backed up by the more tedious serological characters. Clear-cut results
were obtained in which a commercial inoculant strain outcompeted two
The rhizobial component 261
strains isalated from Sudanese sails in accupying nadules an Sudanese
cultivars. Hawever, the calany morphalagy characters used included paly-
saccharide secretian. The inaculant strain produced abundant palysacchar-
ide when grown on yeast-extract-mannital agar, a standard medium for
rhizabia (Bergersen, 1980; Peoples et al., 1989) whereas the twa Sudanese
sail isalates produced little.
It has recently became clear that extracellular palysaccharide (EPS)
productian by rhizobia can be important in successful infection af same
legume roats by rhizobia (reviewed by Brewin, 1991). Of the species
studied, all have infectian thraugh roat hairs but subsequent nodule devel-
opment falls into two types. Thase nodules having an apical meristem
(e.g. lucerne, alfalfa) and consequently of indeterminate growth do not
become invaded by rhizobial mutants lacking EPS. Thase with determinate
growth, such as saybean, can be invaded by mutants lacking EPS. These
twa types af nodulated legumes fall into. different taxanamic groups
(tribes) and their nadules vary in metabalic as well as structural properties
(Sprent, 1980). They are also nadulated by the genera Rhizobium and
Bradyrhizobium respectively. Arachis is not closely related to either of
these ather types af legume: it has a different type af infection pathway
(section 8.2) and a different nadule structure (sectian 8.4). Whether ar nat
EPS malecules are invalved in establishing the symbiasis is not knawn.
Exciting new developments are occurring in unravelling the signal
exchange between plants and rhizobia which leads to successful invasion
thraugh root hairs. One af the first af these important developments was
the discavery that the rhizobial nodD genes can be switched on by the
flavanaid signals from the plant roat (Peters et al., 1986). The nodD genes
have an impartant regulatary role in the aperatian af several af ather nod
genes (Table 8.2). They are also a campanent of the very complex system
which cantrols hast/rhizabial specificity. Even though the infection process
in Arachis appears rather simple, it might be expected that the rhizobial
nod genes wauld still have to. be triggered befare nadulation could pro-
ceed. Evidence that this is indeed the case has been abtained by Smit and
ca-warkers (1990). They used the naw routine procedure of canstructing
genetically engineered rhizobia in which the lacZ gene is inserted in a
pasitian such that it is switched an when nodD is switched an. The lacZ
gene codes far an enzyme, 13 galactasidase, which produces calaured
praducts from suitable substrates. The inductian af nodD can thus be
studied on rhizobial cultures, without the host plant. In this way
Bradyrhizobium sp. NC92, which nodulates Arachis, was campared with
B. japonicum USDA11O, which nodulates soybeans. In both, genistein was
the most active inducer of those tested, but NC92 was 50 times more
sensitive to it than USDA11O. Five inducing compounds were isolated
from cotyledons and seed caats of Arachis: all were active in NC92, none in
USDA11O.
Later on in the development of nodules the precise structure of rhizobial
TABLE 8.2 Nodulation genes of Brady rhizobium (extended from Barbour, Wang
and Stacey, 1992)
nod
A,B,C Involved in infection and early events in nodulation; interchangeable

with relevant genes in Rhizobium, hence called 'common nod genes'.
D May have several copies; involved in regulation and host specificity.
Function unknown.
J Function unknown; membrane based.
K Function unknown.
L,M,N Determine host range in Rhizobium but not confirmed in
Bradyrhizobium.
s, u Confer broad host range in some strains.
v,w Host range.
y Function unknown.
z Host range.
In addition to the nod genes listed, host-genotype specific genes also occur. Most information
is from B. japonicum which nodulates soybean and B. spp. (Parasponia) which nodulates the
non-legume Para5ponia. Available information for B. sp. (cowpea), which nodulates both
Vigna and Arachis, shows that similar genes occur. Symbiotic genes in Bradyrhizobium unlike
those of Rhizobium, are chromosomal.
lipopolysaccharides (LPS) appears important, especially for endocytosis

and bacteroid differentiation. Brewin (1991) suggested that, because deter-
minate nodules undergo endocytosis at a relatively earlier stage and
infected cells then divide, LPS mutants would have an earlier and more
profound effect than in indeterminate nodules where endocytosis occurs
later and infected cells do not subsequently divide. If this suggestion is
correct, it would be expected that the effect of LPS mutants on Arachis
nodule development would be at least as profound as on determinate
nodules such as soybean.
Although most soybean rhizobia do not normally nodulate Arachis,
some can cause production of 'nodule-like structures' (Devine et al., 1983).
Devine and colleagues, in studying host genetics and nodulation, have
isolated a series of rhizobial strains which can nodulate soybeans carrying
the allele rh which normally restricts nodulation. These strains generally
also produce the chlorosis-inducing substance rhizobitoxine. When tested
on two cultivars of Arachis, they produced nodule-like swellings in the axils
of lateral roots but did not produce chlorosis in the shoot. In one case the
swellings were relatively large and rhizobia were isolated from them.
Arachis did not form swellings with any of the non-chlorosis-inducing
soybean strains. In this case the rhizobia are defective in their symbioses
with these two different hosts and it was suggested that they may be fully
compatible with another (unidentified) legume or that they may represent
an active stage in evolution in which specificity is not yet fixed.
Functional nodules 263
The situation is quite different with many cowpea strains of Brady-
rhizobium which can form highly effective nodules on both Arachis
and Vigna unguiculata. Sen and Weaver have studied nodules on these two
hosts in some depth (e.g. 1984) and shown that the nodule structure is
controlled by the host. This overriding effect of the host appears common;
the same rhizobial strain may infect different plants in different ways and
produce nodules with different structures and metabolic processes (Sprent,
1989).
8.4 FUNCTIONAL NODULES
All legume nodules which have been examined assimilate the product of
nitrogen fixation (ammonia) in plant cells using the GS:GOGAT system
(Figure 8.3). The first organic product is thus glutamine. This may be
exported from nodules or further reactions may occur, so that other
compounds are exported, such as asparagine, citrulline, or the ureides
allantoin and allantoic acid (Sprent and Sprent, 1990). The compounds
exported by groundnut have been a matter of controversy for some years.
In addition to asparagine and glutamine, methylene glutamine and ureides
have been reported (review by Schubert, 1986). Several groups have failed
to confirm ureides as a product of N fixation in Arachis. One such study
(Peoples et al., 1986) also resolves the source of 4-methylene glutamine in
the xylem sap of Arachis plants. These workers used nodulated plants,
non-nodulated plants given nitrate and nodulated plants with the root
system purged with Ar02 instead of air. The latter treatment did not affect
nitrogenase (AR) activity but no fixation of nitrogen occurred in the
absence of N2 and thus N compounds in the xylem sap must have arisen
from other metabolic pathways. The overall conclusion was that
4-methylene glutamine is a product of cotyledonary metabolism and its
concentration in xylem sap is unrelated to nitrogen fixation (see also Goto
et al., 1987). A further complication is that unidentified compound(s) in
xylem sap of Arachis species may interfere with the standard ureide assay
(Peoples et al., 1991a). Nitrogen fixing and nitrate assimilating plants
export both glutamine and asparagine, but the amounts vary in a way
which may serve as an indicator of nitrogen fixation (section 8.5).
Legume seeds are well known for their production of lectins - proteins
with pronounced haemagglutination properties. Lectins also figure promi-
nently as a possible constituent of the host-rhizobial recognition system
(Sprent and Sprent, 1990). Strong evidence that they playa part in root
hair infection systems has recently been obtained for white clover (Diaz
et al. ,1989), but this does not appear to be the case for Arachis (Pueppka
et al., 1980), which does not have a root hair infection process and which is
very promiscuous in its relations with different rhizobia. However, lectins
may be involved in nodule development in Arachis. Lectins extracted from
nodules vary from those extracted from seeds: although each organ pro-
duces separate molecules capable of binding to galactose and to mannose,
nodule lectins differ from seed lectins in being glycosylated (i.e. in being
glycoproteins) and, in the case of the mannose binding (ML) one, in amino
acid composition and antigenic proteins (Law et al., 1988). Within nodules,
the ML and galactic/binding lectins (GL) had different distributions
(Figure 8.4), which were interpreted as suggesting a storage function (Law
and van Tonder, 1992). This logical suggestion raises the question as to
why nodules should store proteins. Since Arachis nodules are of determi-
nate growth, storage would not appear to be an insurance against a need
for further growth and yet in earlier work (Kishinevsky et at., 1988) total
lectin concentration in Arachis nodules reached a peak and then declined,
indicating usage or at least breakdown.
Not only are Arachis nodules unusual in their protein storage properties
but also in their lipid metabolism. Seed oleosomes (lipid bodies) have
been known and studied for many years but recent work, mainly from
Bal and co-workers in Newfoundland, has shown that nodules have oIeo-
somes which differ in size and other properties from those of seeds (Table
8.3; Jayaram and Bal, 1991). It was suggested that nodule oleosomes are
transient storage organelles which can be metabolized, whereas seed oleo-
somes are for long-term storage. This suggestion is backed up by studies
with de-topped and darkened plants. Both treatments led to a gradual
decrease in number of oleosomes, coupled with evidence of the 13-
oxidation pathway and glyoxylate cycle in nodule homogenates (Figure
COOH
I
c=o
tH2 From respiration, etc.
I
CH2
I
COOH
2 oxoglutaric acid
NAD+ NADH+H'
GOGAT / " 1 used for further
COOH reactions or exported
COOH
I I
CH.NH2 fH. NH 2
I
2 CH2 2 CH2
I
CH2 CH2
<
I I
COOH CO.NH2
Glutamic acid Glutamine
2NH: )>GS
2 ATP 2 ADP+2 Pi
Figure 8.3 Assimilation of ammonium into glutamine using the enzymes gluta-
mine synthetase (GS) and glutamate synthase, commonly known as GOGAT
(glutamine-2-oxoglutarate-amino-transferase) .
3-celllayer \
contains
Infected tissue tJ 2-celllayer
(no interstitial cells; lacking
large and small galactose-binding
cells mayor not be
oleosomes. lectins
vacuolate;
small oleosomes)
Figure 8.4 Specialized layers in groundnut nodules_
TABLE 8.3 Some differences between nodule and seed derived oleosomes (lipid
bodies) (after Jayaram and Bal, 1991)
Nodule Seed
Approximate % with diameter

< 1 fLm 79 52
1-2 fLm 17 32
> 2 fLm 4 16
% total lipid in certain
fractions C 16:0* 19 12
C 18:0 28 3
C 18:1 14 51
C 18:2 20 29
C 18:3 14 2
Electron dense rim with Present Absent
lipolytic activity
* These values indicate thc number of C atoms, followed by the number of unsaturated
linkages (C=C bonds)
8_5). Compared with cowpea nodules formed with the same strain (Brady-
rhizobium sp. 32H1), groundnut nodules were able to maintain nitroge-
nase (AR) activity for 48 hours after de-topping and darkening, whereas
activity in cowpea fell within hours (Siddique and Bal, 1991)_ Ineffective
nodules accumulated excess lipid (Bal and Siddique, 1991)_ All these data
are consistent with a functional role of oleosomes in nodules.
Glyoxylate cycle
or other metabolic
pathway
Figure 8.5 Simplified pathway for l3-oxidation of fatty acids (AMP, adenosine
monophosphate; ATP, adenosine triphosphate; CoASH, coenzyme A; FAD, flac
Yin adenine dinucleotide; Ppi, pyrophosphate; R, a long chain CH3 (CH2 )n)' The
final product, acetyl CoA, usually enters the glyoxylate cycle in fat-storing seeds,
such as groundnut, and may do the same in nodules. Note the production of
hydrogen peroxide (H2 0 2 ) which may be broken down by catalase (Figure 8.7).
A further point of considerable interest is the location of nodule oleo-

somes. The smaller 'typical' nodule form was largely located in the infected
cells, whereas the larger form, which constitutes only a small fraction of
nodule oleo somes but which is more frequent in seeds, was located mainly
in the three layers of cortical cells immediately exterior to the infected zone
(Jayaram and Bal, 1991). This is the same region (Figure 8.4) in which Law
and van Tonder (1992) found a lack (or only low concentrations) of cells
containing ML in vacuoles. Thus the inner cortical layers appear to be
metabolically distinct from layers exterior to them (as well as from the
infected region - see also Figure 8.2). Distinct layers can also be observed
in other nodules, such as soybean (e.g. Parsons and Day, 1990; James et
at., 1991; Sutherland and Sprent, 1984) and may be related to location of
oxygen diffusion resistance control mechanisms (Figure 8.6). The diffusion
barrier has both fixed and variable components, and enables the supply of
oxygen to be tailored to demand without inactivating nitrogenase; it also
enables the plant to control nodule activity (for example, when under
stress). Barriers have not yet been shown to exist for groundnut but
comparable physiological studies on the latter are under way in our
laboratory.
It has long been known that bacteroids in groundnut nodules are
spherical (e.g. Sen and Weaver, 1984) whereas the same strain of
Bradyrhizobium in cowpea produces rod-shaped bacteroids, which are
Outer cortex
Diffusion barrier
Distribution zone
1%°2
+--- Infected tissue
Figure 8.6 Location of oxygen diffusion barrier in nodules of the soybean type.
very common in legumes generally, though some tribes (Vicieae,

Trifolieae) have nodules with enlarged pleomorphic bacteroids (Sprent,
1980). The spherical bacteroids of Arachis (Figure 8.2) appear to be yet
another apparently unique feature of its nodules. Originally the shape was
thought to be the result of the bacteroid having little or no cell wall
material (to confer shape), so that it adopted a protoplast-like spherical
structure (Staphorst and Strijdom, 1972). More recent studies have shown
that a distinct wall layer is present throughout (Bal et al., 1985). In some
ineffective nodules the bacterial spheres are greatly enlarged (Wilson and
Giller, personal communication). Bacteroids in groundnut nodules occur
singly, each surrounded by a peri bacteroid membrane (PBM) and forming
a peribacteroid unit (PBV), sometimes now termed a symbiosome (e.g.
Brewin, 1991). PBVs containing single bacteroids are found in other
species, such as peas (in which the bacteroids are greatly enlarged and
polymorphic, e.g. Newcomb, 1981), whereas soybeans (for example) may
have eight or more bacteroids per PBV (e.g. Sprent, 1972). The outer
membrane of the PBV is the functional interface between the symbiotic
partners and has many characteristic properties, such as a membrane-
bound ATPase (Brewin, 1991). In the groundnut, the PBM is often very
close to the bacteroid surface and may even touch it. Further, in many
cases where the PBM and bacteroid surface are close or touching, oleo-
somes are found to be close or touching (Bal et al., 1989). In the narrow
peribacteroid space (i.e. between the bacteroid surface and PBM) several
amorphous, electron-dense bodies can be found. These 'dense bodies' (Bal
et al., 1989) remain attached to the bacterial surface after removal of the
PBM and are also seen 'inside' bacteroids, following invagination of the
surface membrane. Often they are seen in intact PBV, near oleosomes. In
these nodules there is a close correlation between nitrogenase and catalase
activities (Figure 8.7). Since dense bodies show a pH-sensitive positive
reaction with diaminobenzidine (DAB), characteristic of catalase, it is
suggested that they are a site of catalase activity. Also showing the DAB
reaction are microbodies in host cells, both near the PBU and towards the
periphery. Since catalase is often observed (e.g. in lipid-storing seeds,
Tolbert, 1981) to be associated with the 13 oxidation pathway (Figure 8.5),
these microbodies and dense bodies may be an integral part of lipid
metabolism in the nitrogen-fixing groundnut nodule.
8.5 MEASUREMENT OF NITROGEN FIX ATION IN THE FIELD
From a practical point of view, farmers and agronomists need to know the
value of nitrogen fixation to the production of an economic yield of grain.
In order to assess this, they require a reliable way of measuring nitrogen
fixation in the field. Numerous techniques have been tried, with varying
degrees of success, and this section will discuss some possibilities.
8.5.1 Acetylene reduction

Although greeted with much enthusiasm and used with success in many
experiments, the acetylene reduction assay has recently been shown to
................. \
10 / \ 50
/
/ \
X" \
/ \
/ \
\
/ \
.,
:; /
/ \
\
\
40
c
"0 / '\
~a.
0 /
c \
0> \
.E 0>
.<:::
'<l-
30 .E
:r: .~
c
U'" .,:J
(5
E '"
c
[(
20
~
u
«
10
Weeks after inoculation
Figure 8.7 Pattern of nitrogenase (acetylene-reducing) activity in groundnut nod-

ules in relation to that of catalase (Figure 8.5). The data give catalase activity in
bacteroids; similar patterns, but at lower levels, were found for host cells. (After
Bal et al., 1989.)
Measurement of nitrogen fixation in the field 269
have many problems as an accurate estimator of N2 fixed (e.g. Witty et ai.,
1986). In many species it has been shown to be reliable only for pot-grown
plants subjected to continuous-flow analytical techniques. Clearly this
precludes assay of field-grown material. The major problems are that many
factors, from plant disturbance to acetylene itself, may result in a decline in
nitrogenase activity. However, species studied in sufficient detail have so
far been few and have not included groundnut. In view of the unique
structure of its nodules, and the fact that nitrogenase activity is retained far
longer after drastic treatments such as de-topping than in other species
such as cowpea (Siddique and Bal, 1991), groundnut may be a species that
is less sensitive to manipulation than other grain legumes. Until more
studies have been carried out it would be unwise to use acetylene reduction
techniques for anything other than a general guide to nitrogenase activity.
Even if it were quantitative, activity is only measured for a short period
(usually minutes) and so there remains the problem of integration over a
season.
8.5.2 Xylem sap analysis

This last objection can also be levelled at assays based on xylem sap
analysis. On the other hand 'instant' measurements are useful for examin-
ing seasonal and diurnal patterns of nodule activity. It is important to
know, for example, which host/rhizobial combinations commence activity
early in plant growth and which persist into the grain-filling period. Are
xylem sap methods therefore any more reliable than acetylene reduction?
As discussed earlier, there are no unique products of nitrogen fixation in
groundnuts, unlike soybeans and related plants where ureides are exported
from nodules. In most agricultural systems, nitrate is the most abundant
form of mineral nitrogen in soil. Nitrate is absorbed by roots and may be
assimilated there or passed to the shoot for assimilation (Figure 8.1). When
the latter occurs, nitrate can be extracted from xylem sap. In a detailed
study of xylem sap composition of cv. Virginia Bunch, in which Nrfixation
was assayed independently by isotope dilution (a method described next),
Peoples et ai. (1986) found that, as plants derived less of their N 2:
(1) the concentration of an asparagine in sap fell;

(2) the ratio of asparagine:nitrate fell; and
(3) the ratio of nitrate:total amino acid rose.
These data suggest that xylem sap analysis could be a valid assay for
nitrogen fixation in Arachis. The method has been tested on field-grown
groundnuts in Malaysia. Results from percentage asparagine in sap and
percentage nitrate in sap are similar (Norhayati et ai., 1988), mean values
being given in Table 8.4. In this work, inoculation improved N2 fixation but
N fertilizer depressed it.
TABLE 8.4 Possible use of xylem sap analysis to estimate nitrogen fixation in the
field
% total Estimated % plant N derived from

xylem sap as N2 fixation using
Site Asparagine Nitrate

(1) (2) (1) (2)
A 71 2 96 90
B 66 1 88 95
C 57 9 67 73
D 53 14 57 65
E 41 30 33 40
Data are from 5 field sites in Malaysia, as reported by Norhayati et ai, (1988), interpreted
using solute data of Peoples et a/' (1986), Site A was inoculated and site E given N fertilizer:
sites B, C and D were neither inoculated nor fertilized,
8.5.3 Isotopes
It is more difficult to integrate N2 fixation over longer periods - up to a
whole season. The simplest case is where soils contain only trace amounts
of combined nitrogen, in which case classic N analysis (usually by Kjeldahl
digestion) may suffice. However, such conditions are relatively rare, The
only reasonable alternatives involve use of isotopes of nitrogen. Of these
13N is radioactive, but has a half-life of about 11 minutes, making it
completely unsuitable for field measurements (and indeed many laboratory
studies). The stable isotope 15N has been the subject of much investigation
and forms the basis of various techniques, two of which have been used to
good effect with groundnut. More details can be found in Bergersen (1980)
and Peoples et al. (1989).
8.5.4 The lSN-dilution technique

Basically this technique involves the incorporation of lsN-labelled sub-
strate into soil and then the growing of fixing and non-fixing plants in this
soil. NTfixing plants will take up some soil 15N but the non-fixing plants
will not use atmospheric N2 (largely 14N). Thus the dilution of lsN in the
fixing plants compared with the non-fixing ones can be used as a measure of
N2 fixation. There are many experimental problems with this technique,
especially in the field - for example, proper incorporation of 15N into all
layers of soil to which roots have access, slow release of lsN from insoluble
to soluble forms during the growing season, and choice of a suitable
'reference' non-fixing plant (Witty, 1983).
Where non-nodulating genotypes are available, these form good refer-
ence plants: they are more likely to have similar rooting behaviour as their
Measurement of nitrogen fixation in the field 271
TABLE 8.5 Use of the 15N dilution technique, using a non-nodulating mutant as a
reference plant, to estimate N2 fixation by groundnuts in the field in India
(after Giller et al. 1987)
Type Genotype Total N in Atom % Estimated

plants kg/ha excess 15N % N fixed
Virginia Robut 33-1 162 0.024 92

NC-17 150
ICGS-22 127 0.023 89
Spanish Gargapuri 131 0.032 89
1-11 111 0.029 90
Valencia PI-259747 126 0.039 86
Non-nodulating
mutant 41 0.039 0
Standard error 4.5 0.0028 1.0
Nodulated plants were grown in soil with 10 kg/ha (NH4hS04 containing 5.123 atom %
excess 15N; non-nodulated plants received 100 kg N/ha at 0.512 atom % excess 15N.
fixing relatives than the different genera often (inevitably) used. Giller et
al. (1987) used a non-nodulating genotype as a reference plant to assess
nitrogen fixation by seven groundnut genotypes, grown in an Indian alfisol
which had previously been used for Zea mays to reduce soil N to a low
level. Some of the data are shown in Table 8.5. It can be seen that the
amount of N fixed varied from 100 to 152 kg/ha over the 89-day growing
season, and that the virginia types generally fixed more than the valencia or
spanish types. The percentage of N fixed by the different genotypes did not
vary greatly. Recovery of soil N by the non-nodulating genotype was poor.
These data not only show the utility of the 15N dilution technique but also
suggest that total plant N is a good way of comparing N-fixing potential of
different ground nut genotypes.
The superior performance of virginia types has been shown elsewhere.
For example, Arunachalam et al. (1984) looked at 19 characters in 21
genotypes in India. Of these genotypes, six were virginia bunch and five
virginia runner types - and they all performed better for most characters
than the five spanish and five valencia types. Nodule mass, estimated at the
S2 stage of growth was found to be a good indicator of nitrogen fixing
potential in this case, as was acetylene reduction activity measured in the
'old-fashioned' closed system assay.
8.5.5 'Natural abundance' 15N method

This method has become more common during the last few years, based on
the fact that (usually) soil N contains more 15N than atmospheric N2 . Thus
the difference (8 15N) in 15N between plants reliant on soil N (or on the
proportion of 15N in soil itself) and those fixing N2 can be used as an
estimate of N2 fixation. The method has been well discussed by Shearer
and Kohl (1986). It is most useful where the 15N content of soil is fairly
uniform and where the difference between 15N content of soil and atmos-
phere is greatest. It avoids the problem of incorporation of 15N into soil but
needs much more careful analysis and a more sensitive mass spectrometer
than the isotope dilution technique. It may be the only feasible method for
deep-rooted plants, especially trees, but has also been applied successfully
to grain legumes.
For groundnut, values from 25% to 64% of plant N derived from
fixation were quoted by Peoples et al. (1991b). The data given were from a
variety of experiments involving different cultivars, crop rotations and
levels of water supply. N2 fixation generally supplied a lower proportion of
plant N than in other studies quoted, but the higher proportions were
correlated with higher amounts of N2 fixed. Thus the low values may
reflect the fact that treatments inhibited N2 fixation more than assimilation
of soil N, a common phenomenon (Sprent et al., 1983).
8.6 EFFECTS OF ENVIRONMENTAL STRESS IN NODULATION

AND NITROGEN FIXATION
Compared with those for other grain legumes, studies on the effects of
environmental variables on nodulation and' N2 fixation by groundnuts are
comparatively rare and comparison of these effects with utilization of soil
N even more so. However, there are several reports which deal with the
direct interactions between soil Nand N2 fixation. Although groundnuts
can take up and use mineral nitrogen (as already explained), there is
evidence that they do not do so as readily as some other grain legumes,
such as soybean. For example, Nambiar et al.(1986) found that groundnut
was a poor user of fertilizer N when compared with sorghum. They
examined nitrate reductase activity and found that it was low in leaves of
both nodulating and non-nodulating genotypes, even when leaf nitrate
content was high. This leaf nitrate may have been in a non-metabolic pool
(vacuole) as has been found in various plant species. These workers found
that nitrogenase activity declined during pod-fill, but that addition. of
mineral N at this stage was without effect. Their results are consistent with
those of Sung and Sun (1990), who found that nitrate reductase activity
fell during pod fill but who also found that nitrogenase activity (AR)
increased during pod-fill. Both genotypic and environmental effects may
explain this discrepancy in nitrogen fixation during pod-fill but the results
do suggest that selection for better N2 fixation may be worthwhile.
It has been known for many years and in several agricultural legumes
that nitrate affects root hair infection and nodule function (Sprent and
Environmental stress in nodulation and N fixation 273
Sprent, 1990) but it is not known how the infection pathway found in
groundnut responds, or how nitrate affects nodule functioning. In several
species it has been shown that nitrate increases the oxygen diffusion
resistance in the cortex of nodules and in this way inhibits (reversibly,
within limits) N2 fixation. However, as seen above, groundnut nodules
have cortices and infected regions with characteristic structural features:
these may respond differently to nitrate and other forms of stress. Detailed
studies on this are urgently required.
8.6.1 Water
Water stress is often a problem in areas where groundnut is grown (Chap-
ter 9). There is evidence that groundnut nodules are less sensitive than
those of some other legumes to soil water deficit. Venkateswarlu et al.
(1989) compared the responses of ground nut and cowpea. In cowpea,
nitrogenase activity ceased when leaf water potential fell below -0.9 MPa
whereas the corresponding value in ground nut was -1.7 MPa. Nodules on
cowpea, but not on groundnut, were shed under stress. In the latter, stress
resulted in a sharp increase in total soluble sugars and leghaemoglobin.
Devries et al. (1989), comparing groundnut with soybean and pigeon pea,
found that groundnut alone responded to drought by increasing its pro-
portion of roots below 30 cm. These workers also found groundnut nitroge-
nase activity to be continued longer into the drought period than that of the
other legumes and coupled this with a higher leaf water potential and
stomatal conductance. The comparative drought tolerance of groundnut
thus seems to result from both nodule and plant factors.
8.6.2 Iron and bicarbonate

Groundnuts are very susceptible to lime-induced iron deficiency (Chen and
Barak, 1982). Legume nodules have a particularly high requirement for
iron, because of its incorporation into both nitrogenase and haemoglobin.
Iron-deficient soils in which groundnuts are grown are often alkaline, with
a high bicarbonate content. Recent work has shown that both iron and high
or low pH can separately affect nodulation and nitrogen fixation (O'Hara
et al., 1988a,b; Tang et al., 1991). Effects are complex and involve both
host and rhizobia. In common with many soil micro-organisms, bradyrhi-
zobia can secrete siderophores to aid Fe uptake. However, strains vary in
their ability to do this (O'Hara et al., 1988b). One strain which produces
siderophores, NC92, was found to nodulate groundnuts better in low Fe
conditions than strain TAL 1000, which does not (Tang et al., 1991).
Nodulation, nodule development and nodule functioning were all reduced
at low iron concentrations. As with most legumes, bicarbonate adversely
affected nodulation and nodule functioning and effects were more pro-
nounced on NTfixing plants than on those given mineral N. Table 8.6 gives
some examples of the effects of Fe and bicarbonate on nodule parameters,
taken from Tang et ai., (1991).
8.6.3 Salinity
Another soil factor often encountered by groundnut is salinity. This may
affect nodule initiation, although interestingly NaCl causes normally bare
roots to produce hairs (Sprent and McInroy, 1984). Arutyunova and
Shevyakova (1984) looked at the effects of both NaCI and Na2S04 on
growth of nodulated groundnuts fed with NHt and NO). However, the
work concentrated on mineral N and it was found that ammonium could
partially offset the adverse effects of salinity. On plants fed with am-
monium, nodules were formed in the presence of 34 mol/m 3 of NaCl, but
14 mol/m 3 for Na2S04' The reverse was true for nitrate-grown plants.
These data show clearly that the effects of salinity on nodulation are
complex but little light is shed on the mechanism of its action.
8.6.4 Phosphate
In many soils, nodulation is limited by available phosphate, although not
all legumes are equally sensitive. Possession of endomycorrhizas generally
TABLE 8.6 Effect of iron on nodulation and nitrogen fixation by groundnut cv.
Tainar, inoculated with strain NC92 (after Tang et aI., 1991)
Age at Concentrations Number Plant Nin

harvest of Fe of nodules/ dry weight shoot
(days) (mmollm 3 ) plant (g) (mg)
20 0.05 14 0.42
0.4 26 0.40
7.5 40 0.49
7.5 + HCO) 2 0.41
30 0.05 133 0.83 12
0.4 208 0.97 16
7.5 202 1.07 15
7.5 + HCO) 26' 1.01 11
50 0.05 172 0.97 16
0.4 374 2.03 40
7.5 389 2.57 49
7.5 + HCO) 101 1.33 19
Plants were grown in solution culture pH 5.5 in a greenhouse. In one treatment bicarbonate
was given by including CaC0 3 and NaHC0 3 , each at 5 mol/m 3 : the pH of this solution was
8.6-8.7.
Prospects for improvement 275
TABLE 8.7 Effect of inoculation with the mycorrhizal fungus Glomus clarum on
nodulation and shoot dry weight of 5 genotypes of groundnut grown in sand
containing rock phosphate for 90 days
Shoot D.W. g
Nodule rating
Type Genotype (+MrM) x 100 -M +M
Virginia Robut 33-1 138 1.54 2.18

leGS 15 131 1.74 2.24
Spanish Gangapuri 156 1.13 1.60
J-11 145 1.25 1.86
Valencia PI 1259747 147 1.78 2.41
+ M: mycorrhizal plants
-M: non-mycorrhizal plants
Note that (as in Table 8.4) the spanish types performed less well.
aids uptake of P from relatively insoluble forms such as rock phosphate.

This is certainly true for ground nut. In a detailed study of several geno-
types, Daft (1991) found that mycorrhizal infection increased nodulation
and shoot dry weight of plants grown on rock phosphate from a variety of
sources. However, genotypes differed markedly in response: increases of
shoot dry weight varied from 40% to 360%. Detailed inspection of the
nodulation data showed far more variation in the absence than in the
presence of mycorrhizal infection (Table 8.7). These interesting data
suggest that assessment of nodulation potential might best be carried out
on mycorrhizal plants. In low-P soils, plants are likely to be naturally
mycorrhizal.
8.7 PROSPECTS FOR IMPROVEMENT
Many possibilities have been touched on in earlier sections. Selection of

both host and rhizobial genotypes can give improved NTfixation in labora-
tory experiments. However, one problem with nodulation in the field is
competition from indigenous rhizobia, and this has been studied at
ICRISAT over many years. Nambiar et al. (1984) give data for various
inoculant strains, host cultivars, seasons (rainy, dry) and methods of
inoculation. Not surprisingly, a range of responses was found, but there
were consistent increases in economic yield with cultivar Robut 33-1 given
inoculant strain NC92.
There are various practical problems associated with inoculation. In the
study quoted above, liquid inoculants were sometimes found to be better
than granular forms, but this depended greatly on soil moisture. Further,
these workers point out that groundnut seed may be too fragile for seed
inoculation (as generally used for soybeans, for example) and fungicide
treatments may be toxic to rhizobia. Other management practices which
may affect nodulation include weed control. In two seasons of field trials in
India, Malavia and Patel (1989) found that pre-emergence application of
three weed-control chemicals adversely affected groundnut nodulation.
Taking improvements in a more general sense to include soil fertility,
can groundnuts in a crop rotation significantly reduce N-fertilizer require-
ments for non-fixing crops? This is a complex question for all legumes,
especially grain forms, where much of the N is removed with crop. If more
N is removed than fixed, then soil N will be depleted and this sometimes
occurs. Further, the C:N ratio of legume residues (as with any plant
residues) will affect how quickly N is mineralized and made available to
subsequent crops (Sprent, 1987). With these cautions in mind, there is
evidence that groundnuts can have a positive effect on soil N in subsequent
crops. Dakora et al. (1987) compared groundnut and cowpea in the field in
northern Ghana. Both legumes nodulated well with indigenous rhizobia.
Groundnut fixed 101 kg N/ha (estimated by difference in N content of
groundnut and maize grown in the same soil), of which 46% was removed
with grain. The corresponding values for cowpea were 201 kg/ha and 34%
removed. Thus cowpea had far more N in its residue than groundnut
(150 compared with 68 kg/ha). In spite of this the two legumes had approxi-
mately equal effects on a subsequent maize crop, equivalent to 60 kg N/ha.
In other words a larger proportion (60%) of N in the groundnut residue
was made available to the subsequent maize crop than cowpea residue
(27% ).
ACKNOWLEDGEMENTS
I should like to thank Dr M.B. Peoples and Dr I. Law for permission to

quote data in advance of publication. Production of this chapter relied on
the technical skills of Ms M Gruber (photographs) and Mrs I Pimbley
(diagrams) .
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CHAPTER 9
Groundnut water relations

G. C. Wright and R. C. Nageswara Rao
9.1 INTRODUCTION
About 80% of the world groundnut production comes from seasonally

rainfed areas in the semi-tropics, where climate is characterized by the
low and erratic rainfall. Drought is recognized as one of the major
constraints limiting groundnut productivity in these regions (Gibbons,
1980). This chapter reports various aspects of groundnut water relations,
including the effect of water deficits on a range of physiological and
morphological processes which affect shoot and root growth, and ulti-
mately pod yield. Extensive reviews on this subject have been published
over the past decade (for example, Boote et al., 1982; Boote and
Ketring, 1990). The topics discussed in this chapter are intended to sup-
plement and update, where necessary, the literature reviewed in these
excellent papers. These reviews concentrate particularly on groundnut
irrigation management and so readers are referred to them for detailed
information in that area. This chapter therefore concentrates more on
potential genetic and management solutions to the problem of drought in
groundnut.
9.2 GROUNDNUT WATER STATUS
Leaf water status affects numerous physiological processes which contrib-

ute to plant growth and yield. The status of water in plants represents an
integration of atmospheric demand, soil water potential, rooting density
and distribution, and is therefore a true measure of drought stress in plants
(Kramer, 1969). The water status of a crop plant is usually defined in terms
of its water content, water potential, or its components, viz osmotic and
turgor potential (Turner, 1986a).
282 Groundnut water relations
9.2.1 Water content
Leaf relative water content (RWC) has been successfully used to monitor
water content and status in groundnuts (e.g. Bennett et af., 1981, 1984;
Slatyer, 1955). Sinclair and Ludlow (1985) argue that RWC is a more
useful integrator of plant water balance than leaf water potential and
should provide universal relationships between physiological traits and
level of drought stress. RWC values in well-watered groundnuts are typi-
cally in the range of 85-98% (Bhagsari et af., 1976; Joshi et af., 1988;
Bennett et af., 1981, 1984; Prabowo et af., 1990) Under drought con-
ditions, RWC as low as 29% has been measured (Bhagsari et af., 1976),
indicating that ground nut has a very low lethal water status, This attribute
should contribute to high levels of dehydration tolerance and leaf survival
in groundnut during intermittent drought stress (Ludlow and Muchow,
1988), in a similar fashion to that reported for pigeon pea (Flower and
Ludlow, 1986).
Care needs to be taken when measuring RWC in severely stressed
groundnut leaves. Wright (unpublished data) found that the standard 4 h
floating time, recommended by Barrs and Weatherley (1962) and Bennett
et af. (1984), was insufficient for the full rehydration of severely stressed
groundnut leaves. For these leaves a floating time of between 8 and 12 h
was required to achieve full turgidity. Thus, errors in RWC of up to 25%
were recorded when 4 rather than 12 h floating time was provided.
9.2.2 Plant water potential

The establishment of plant water status on a sound thermodynamic basis
by the introduction of the concept of water potential has led to the
widespread adoption of total water potential as a measure of plant water
status (Slatyer and Taylor, 1960). The total water potential, usually in
leaves because of ease of measurement, can be partitioned into the major
components of osmotic and turgor potential (Turner, 1986a). In ground-
nut, problems with phloem sap exudation can make determination of the
end-point difficult when the pressure chamber technique is used (Pallas et
af., 1979; Bennett et af., 1981, 1984). This problem has limited leaf water
potential research in groundnut (Bennett et af., 1981) and led to more
widespread use of thermocouple psychrometers for leaf water potential
measurement. Wright et af. (1988b) compared thermocouple psychrometer
and pressure chamber techniques and concluded that the pressure chamber
over-estimated leaf water potential in ground nut by about 0.4 MPa. This
error can lead to large errors in leaf turgor potential, where the pressure
chamber is used for measurement of total potential component. It was
suggested, however, that the pressure chamber technique may be appro-
priate in comparative studies of groundnut water status. A procedure was
described to aid end-point identification when using groundnut leaves in
Groundnut water status 283
the pressure chamber. Rajendrudu et al. (1983) also reported that the
hydraulic press was suitable for leaf water potential measurement in
groundnut.
Leaf water potential (tjJl) measured about midday in field-grown ground-
nuts under well-watered conditions has been reported to range widely from
about -0.6 to - 2.85 MPa (Joshi et al., 1988; Pallas et al., 1979; Patel et al.,
1983; Bennett et at., 1984; Allen et al., 1976; Erickson and Ketring, 1985;
Stone et al., 1985). The lower (more negative) values of tjJl measured under
well-watered conditions (Stone et al., 1985; Erickson and Ketring, 1985)
occurred in semi-arid regions where plants were presumably under some
environmental stress. Ong et al. (] 985, 1987) have shown that tjJl decreases
linearly with increasing vapour pressure deficit. '
In droughted crops, tjJl in groundnut decreases markedly in response to
declining soil water availability and atmospheric demand. Values in the
range of -3.0 to -4.5 MPa have been reported in drought stressed crops in
the field (Bennett et al., 1984; Joshi et at., 1988; Pallas et al., 1979; Stone et
at., 1985; Erickson and Ketring, 1985), while some reports suggest that
groundnut can reduce its tjJl to even lower values such as -6.3 MPa
(Sarma, 1984), -8.3 MPa (Allen et al., 1976) and -10.0 MPa (Ludlow
and Muchow, 1988).
While transpiration rate and related processes are closely coupled to tjJl,
it has become evident in recent times that physiological and morphological
growth processes are more closely coupled with the osmotic (tjJ1T) and
turgor (P) pressures of tissues (Begg and Turner, 1976; Turner, 1986a). In
particular it is believed that P is the transducer of water deficits in the cell
(Hsiao, 1973) and physiological and morphological processes such as
stomatal closure and leaf rolling occur at zero P (Turner, 1974; Wright et
al., 1983b; Ludlow et at., 1985). There is also recent evidence that the root
acts as a sensor of plant water deficit, possibly through turgor pressure
regulation in the growing root (Turner, 1986a).
Turgor pressure is usually calculated as the difference between tjJl and
tjJ1T. In groundnut, relationships among water potential components have
been measured and indicate that zero P occurs at tjJl of between -1.3 and
- 2.0 MPa in field (Bennett et al., 1981, 1984; Erickson and Ketring, 1985)
and controlled environment (Black et al., 1985) studies. Variation in the tjJl
at which zero P occurs is indicative of osmotic adjustment, a common
adaptive response which allows turgor maintenance through active solute
accumulation and lowering in tjJ1T (Begg and Turner, 1976). There are,
however, conflicting reports concerning the extent of osmotic adjustment
in groundnut. Bennett et al. (1981, ]984) reported it was small in cv.
Florunner, although it was suggested that some adjustment in the order of
0.3 to 0.6 MPa occurred in response to midday tissue water deficits and
naturally occurring soil water deficits. Similarly, Black et al. (1985) found a
small degree of adjustment (0.3 to 0.4 MPa) in cv. Robut 33-1, while
Ong et al. (1985), in a similar controlled environment facility, found no
significant long-term adjustment. On the other hand, Erickson and Ketring
(1985) reported substantial osmotic adjustment in field-grown groundnut,
ranging from 0.6 to 0.9 MPa. There was also evidence that there were
significant cultivar differences in the extent of adjustment. Stirling et ai.
(1989b) showed that, while substantial osmotic adjustment between 0.84
and 1.58 MPa occurred in expanding leaves, it was virtually non-existent in
mature leaves. This response allowed expanding leaves to maintain higher
turgor levels during periods of stress. Interestingly, turgor remained high
in pegs despite severe plant water deficits experienced during early and late
reproductive development. The situation regarding the importance, extent
and possible cultivar variation in osmotic adjustment in groundnut is
therefore unclear. Further research is needed to assess the influence of
other factors, such as severity of stress and radiation levels, on the ex-
pression of osmotic adjustment in groundnut.
9.3 EFFECT OF WATER DEFICIT ON SOME PHYSIOLOGICAL

AND MORPHOLOGICAL PROCESSES
9.3.1 Photosynthesis and stomatal conductance

Bhagsari et ai. (1976) found that net photosynthesis (Pn) in potted ground-
nut plants was not affected by increasing stress until shortly before wilting
commenced. Pn had declined to between 8% and 13% of well-watered
controls (maximum Pn rate 37.7 mg CO 2 /dm/h) by the time IjJI reached
-3.0 MPa. Their data indicated the major effect of drought on Pn was
exerted through stomatal closure and suggested that stressed plants stoma-
tal conductance (g/) is often higher than is necessary to satisfy photosynthe-
tic CO 2 requirements. Similarly, Pallas and Samish (1974) and Pallas et ai.
(1974) reported parallel diurnal trends in Pn and transpiration which were
apparently due to stomatal effects.
Unfortunately there is a lack of published information on the influence
of water deficits on photosynthesis in groundnut grown under field con-
ditions. The situation with regard to g/ is somewhat better, no doubt due to
the lower cost and availability of porometers compared with portable
photosynthesis chambers. Sivakumar and Sarma (1986) made diurnal
measurements of g/ in groundnut on a range of water stress treatments in
Central India. They showed that both the time of day and the degree of
drought stress influenced g/. In moderately stressed plants g/ increased until
1300 h, followed by a dramatic decline in the afternoon. In severely
droughted plants, following relatively high g/ in the early morning, stoma-
tal closure occurred by 1100 h in response to reduced soil water avail-
ability. Such a stomatal response, which enables stomata to open briefly
and photosynthesise during the morning when evaporative demand is
low and to close stomata when demand increases later in the day, is an
Deficit effects on physiological/morphological processes 285
important adaptive feature. Similar stomatal adaptations in response to
water deficits have been reported for other crops (Turner, 1974; Turner et
ai., 1978).
Maximum stomatal conductance in groundnut of about 3-4 cmls under
well watered conditions, and of greater than 2 cmls in wilted leaves are
exceptionally high compared with many temperate species (Black and
Squire, 1979; Black et ai, 1985; Devries et ai., 1989a). The rapid and almost
complete recovery to high stomatal conductance following severe stress has
been widely reported in groundnut (Allen et ai., 1976; Pallas et ai., 1979;
Black et ai., 1985; Joshi et ai .,1988). Presumably plant water status and
leaf photosynthesis also return to near normal following severe stress. This
capacity to recover fully from stress represents an important adaptive
response in groundnut.
Bennett et ai. (1984) showed that stomatal conductance in groundnut
was curvilinearly related to plant water status, with g, declining rapidly
below a tVl of -1.4 MPa and a P of 0.1 MPa. Other workers report that g,
declines linearly with tVl (Stone et ai., 1985; Black et ai., 1985) and that g, is
poorly correlated with bulk leaf P. The ability to maintain high g, at low IjIl,
known as stomatal adjustment, is a common adaptive response to drought
(Ludlow, 1980). Such adjustment has been observed in groundnut (Black
et ai., 1985) and within groundnut cu]tivars (Joshi et ai., 1988); however,
the physiological mechanisms responsible for this adaptation are unknown.
Black et ai. (1985) argue that osmotic adjustment, which can maintain high
turgor pressure around the stomatal cavities in other species (Brown et ai.,
1976; Wright et ai., 1983b; Turner et ai., 1978), is not responsible for
stomatal adjustment in groundnut.
Vapour pressure deficit (VPD) has also been shown to influence g, and
photosynthesis in groundnut (Black and Squire, 1979; Tsuno, 1975; Stone
et ai., 1985; Erickson et ai., 1986; Azam-Ali, 1984). Radiation-use
efficiency (RUE, dry matter production per unit of light intercepted) in
groundnut declines in response to increasing VPD, largely due to stomatal
closure arising from lowered IjII at high VPD (Ong et ai., 1987).
9.3.2 Leaf area development and display

A reduction in water use in groundnut can be achieved by limiting leaf area
development and via paraheliotropic leaf movements, in addition to that
achieved through stomatal control (mentioned above). Regulation of
water use through reductions in leaf area development, or changes in leaf
orientation, rather than through reductions in g, is possible when drought is
imposed slowly. In this situation high g, permits high assimilation rates to
be sustained in the remaining leaves for extended periods of water deficits
(Black et at., 1985).
The reduction in leaf area development in groundnut in response to
water deficits can arise from reduced rates of leaf initiation (Ochs and
Wormer, 1959; Billaz and Ochs, 1961; Boote and Hammond, 1981; Ong
et al., 1985) and reduced rates of leaf expansion (Pandey et al., 1984b; Ong
et al., 1985; Ong 1986). Leong and Ong (1983) found that initiation of
leaves was more sensitive to mild water deficits than were flower, peg or
pod initiation. Leaf number has been shown to be greatly reduced by stress
(Ong et al., 1985). A linear decline in the relationship between the thermal
time to produce one leaf and IjJI below -0.6 MPa was shown by Harris et al.
(1988). This relationship was similar for four cultivars used in this study.
Soil water deficit inhibits leaf expansion through a reduction of RWC
(Slatyer, 1955; Allen et al., 1976) or leaf turgor potential (Ong et al., 1985).
In the absence of temperature limitations, the rate of leaf extension (and
hence expansion) was linearly related to P in the range of 1.0 to 0 MPa, or
as IjJI declined from -0.5 to -1.5MPa (Ong, 1984; Ong etal., 1985).
Variation in the water content of leaves was the main factor controlling P,
since diurnal changes in 1jJ7r were small, and no significant long-term
osmotic adjustment occurred in expanding leaves. Black et al. (1985)
suggests that persistent small differences in water status may therefore
induce large changes in final leaf size, since the duration of expansion is
almost unaffected by water stress. Assimilate shortages during the night
can further reduce leaf-expansion in water-stressed groundnut leaves (Ong
et al., 1985).
Leaf adaptations that change leaf angle and orientation decrease the
radiation load on groundnut leaves. Such movements reduce leaf tempera-
ture and the vapour pressure gradient at a time when stomata are nearly
closed and leaves are least able to lose heat by transpirational cooling
(Gates, 1968). Mathews et al. (1988b) have shown that the degree of leaf
movement was influenced by IjJI, with significant movement (as measured
by the ratio of fractional radiation interception to the square root of leaf
area index) occurring between -0.8 and -1.2 MPa. The observations that
stomatal conductance and leaf expansion cease at similar IjJI levels in the
field (Bennett et al., 1984; Ong et al., 1985) suggests that all of these
processes provide an adaptive capability for optimizing water loss and
restricting leaf temperature over a wide range of soil water deficits. Boote
et al. (1982) give a more comprehensive review of the literature on leaf
display, and description of likely mechanisms responsible for parahelio-
tropic movements in groundnut.
Leaf senescence, or abscission, is often considered to be a mechanism
for conserving water during periods of soil water deficits. Leaf senescence
can occur in groundnut, particularly when exposed to late season drought
(Pandey et al., 1984b), but the crop is considerably more able to retain leaf
area despite severe soil water deficits than many other grain legumes
(Pandey et al., 1984b; Devries et al., 1989a). Few attempts have been made
to quantify leaf senescence in groundnut, although Boote et al. (1986)
included a leaf senescence function in the PNUTGRO model. Leaf senes-
cence is an irreversible process; and, although it is often considered a
Effect of water deficits on root and shoot growth 287
mechanism for conserving water during dry periods, it constitutes a loss of
potential production if the stress is subsequently relieved.
9.3.3 Nitrogen fixation

In groundnut, nitrogen (N) is vitally important to the process of leaf
photosynthesis (Hubick, 1990), radiation use efficiency and biomass
accumulation (Wright et al., 1993b). Information on the influence of water
deficits on nodulation and N fixation in groundnut is limited (Boote and
Ketring, 1990). Recent data has demonstrated that N fixation is extremely
sensitive to periods of soil water deficits (Venkateswarlu et al., 1989) and is
reduced weII before visible stress symptoms ~ppear (Devries et al., 1986;
1989a). Nitrogenase activity has been shown to be severely reduced as leaf
and nodule tV decrease below -1.4 MPa (Devries, 1989a). Little is known
about the relative sensitivity of N fixation and photosynthesis in ground-
nut. In soybean, the effects of soil dehydration on N fixation occur at much
higher fractions of transpirable soil water than do the effects on the carbon
accumulation processes (Sinclair, 1986; Sinclair and Ludlow, 1986), indi-
cating that this response may be widespread in legumes. Clearly, greater
sensitivity to soil water deficits by N accumulation compared with carbon
accumulation could result in greater yield reductions than those expected
based on carbon limitation to drought alone. More research on ground nut
productivity and N accumulation in response to drought is clearly
warranted.
9.4 EFFECT OF WATER DEFICITS ON ROOT AND SHOOT

GROWTH
9.4.1 Root growth and soil water extraction

The extent and function of roots are influenced by seasonal, genetic and
environmental factors (Klepper, 1987). They are discussed below in re-
lation to observations made in groundnut.
( a) Seasonal factors
The apparent root depth progression (ARDP), estimated by examining the
time taken by a crop to extract soil water from particular soil layers,
provides a method to evaluate seasonal progression of groundnut root
growth. Boote et al. (1982) reported that the ARDP for cv. Florunner
progressed steadily at 2.2-2.8 cm/day until 130 days after planting. In
contrast, ARDP rates in four groundnut cultivars ranged between 1.0 and
1.2 cm/day (Mathews et al., 1988a). Plant population can markedly affect
ARDP, with values ranging from 1.2 to 2.0 cm/day in a subtropical
environment (Wright and Bell, 1992b), and 0.85 to 2.3 cm/day in a semi-
arid environment (Simmonds and Williams, 1989). It is evident that the
rate of root penetration into the soil appears to depend on the soil type,
temperature, cultural practices and the aerial environment (Boote and
Ketring, 1990). For instance, Simmonds and Ong (1987) report that ARDP
was 2.0 compared with 1.5 cm/day for plants exposed to a VPD of 2.5 and
1.0 kPa, respectively.
There are conflicting reports about the seasonal pattern of root growth in
groundnut. In some studies, the proportion of biomass allocated to roots
declines markedly as the season progresses. For instance, while 37% of the
total biomass was contained in roots at 21 days after planting, only 1.5%
was recorded at harvest (McCloud, 1974). Similarly, in the rainy season
crop studied by Gregory and Reddy (1982) total root length did not
increase after 55 days after planting. In contrast, total root length nearly
doubled between c. 60 and 90 days after planting in groundnut crops grown
under droughted conditions (Robertson et al., 1980; Nageswara Rao et al.,
1989b). It appears that groundnut is capable of a large investment in root
growth throughout the season, although Nageswara Rao et al. (1989b)
suggest significant root growth during pod filling may only be maintained
when water is severely limited.
(b) Genetic factors

The superior ability of groundnut to maintain favourable water status
during periods of soil water deficits, in comparison to soybean and pigeon-
pea was related to greater proliferation of roots in deeper regions of the
rooting zone (Devries et al., 1989a). Similarly, Pandey et al. (1984b)
concluded that the higher root densities in groundnut at lower soil depths
conferred superior drought tolerance compared to soybean and mungbean.
Groundnut cultivar differences in a number of rooting characteristics
have been shown to exist in glasshouse and field studies. The extent of the
root depth and root length density becomes very important for soil water
extraction during periods of prolonged water deficits. The extension rate of
the tap root up to 32 days of age, and the rate of descent of the soil water
extraction front for the entire season, varied between four cultivars under a
prolonged drought in the field (Mathews et at., 1988a).
However, only minor differences in patterns of water extraction and total
water use were observed among cultivars. Based on their limited cultivar
range, they concluded that the extent to which root characteristics and water
extraction patterns contribute to yield variation in groundnut was small. In
another field study, Wright et al. (1991) have shown substantial cultivar
variation in water extraction at depth, and total water use, exists in
groundnut. The cultivar Virginia Bunch was able to extract c. 40 mm more
soil water at depths below 70 cm compared to three other cultivars (Figure
9.1). Similarly, Chapman (1989) found that cv. Virginia Bunch extracted
significantly greater amounts of soil water (c. 12-30 mm) than a range of
other cultivars subjected to a drought in the pegging stage on a deep
krasnozem soil. These studies have clearly demonstrated that considerable
variation in water extraction from deep soil layers exists among groundnut
cultivars. Based on these results, it is clear the effort required to develop
large-scale screening methods for breeding programmes may well be justi-
fied. Deep-rootedness and faster extraction may be very appropriate in
tropical environments where groundnut is grown solely on stored moisture
in the dry season (often following rice) on deep and high water-holding
capacity soils (Prabowo et al., 1990).
Substantial cultivar variation in a number of root characteristics of
groundnut has been shown under controlled conditions. Ketring et al.
(1982) and Ketring (1984) found that cultivars differed in root volume, root
dry weight, root length and number. Similar cultivar variation was
measured in 22 cultivars grown in hydroponics, where large differences in
root length, volume and dry weight were apparent (Pandey and Pendleton,
1986). A strong correlation between root dry weight and shoot dry weight
seems to exist in groundnut as evidenced from data reported from glas-
shouse (Ketring, 1984; Pandey and Pendleton, 1986) and field studies
(Wright et al., 1994). Figure 9.2 presents this data, which suggests that
indirect selection for large root systems may be possible under field
conditions.
There have been few quantitative studies showing that the differences in
rooting characteristics mentioned above are important in allowing one
cultivar to perform better than another under a specific drought regime.
Higher yield and improved drought tolerance have been associated with
larger and deeper root systems in wheat (Hurd, 1974), sorghum (Wright
and Smith, 1983) and rice (Steponkus et al., 1980). Presumably a similar
association between root size and yield exists in groundnut, although no
definitive data are available which categoricaUy support this hypothesis.
Considerable research effort is required to show that such an association
exists in groundnut. Simple, rapid and inexpensive methods to screen for
rooting characters which correlate with observed field patterns need to be
developed. For instance, Ketring et al. (1985) and Ketring (1986) report
that 'root-effectiveness' can be quantified by measuring the apparent sap
velocity (Av). Under well-watered conditions in the glasshouse, Av ranged
from 0.8 to 1.2 cm/min, while under stress it declined to less than 0.5 cm/
min. Cultivar variation in Av was apparent, and plant growth measurement
indicated that the greater Av was probably due to differences in root
function rather than root mass.
(c) Environmental factors

The depth and distribution of root length in groundnut is dependent on soil
type (Boote and Ketring, 1990). In sandy soils, groundnut roots have been
Volumentric water content (%)
o 10 20 30 40 50
o
\
\
\
20 o ,,
'0
\
\
40 o
"-
""o
\
60 \
\
o
I
/
/
80 o
Soil I
I
depth I
100 I
(cm) o
I
I
120
140
160
180
Figure 9.1 End-of-season volumetric soil water content profile for four cuItivars
grown under a terminal drought stress: (0) Virginia Bunch; (e) Q18801; (6)
McCubbin; (.) Red Spanish. The initial volumetric soil water content measured 2
weeks after planting is also shown (0 - - - D). (Reproduced with permission, Wright
et at., 1991.)
35
30 • •
•
25 •
• •
C 20
'"
~ •
j
15 •
~
"0
(5
•
•
0
£
en 10
•
•
5
0 ~
0 2 3 4 5 6 7 8
Root dry wt (glplant)
Figure 9.2 Relationship between root dry matter and shoot dry matter in a
number of glasshouse and field experiments. Data from: (0) Ketring, 1984; (-)
Pandey et at., 1984b; (e) Wright et at., 1992.
measured to a depth of 150 cm (Robertson et at., 1980) and greater

(Hammond et at., 1978). On loam and clay soils, roots and soil water
extraction have been recorded at depths in excess of 100 cm (Nageswara
Rao et at., 1989b; Mathews et at., 1988a; Prabowo et at., 1990; Wright et
at., 1991). Ludlow and Muchow (1988) warn that measurements of rooting
depth and root length density in crops do not necessarily give an indication
of a genotype's ability to extract soil water. Measurements of soil water
extraction can, however, provide useful indirect information on root func-
tion (Ketring, 1986).
Soil water content can affect root depth and distribution in groundnut.
As surface layers dry out, the pattern of water extraction shifts and a
proportionately smaller part of the root system in the subsoil is responsible
for water uptake (Mantell and Goldin, 1964; Allen et at., 1976). Root
length density profiles reported by Devries et at. (1989a) and Pandey et at.
(1984b) are shown in Figure 9.3, and illustrate how drought increased root
length density in the lower profile compared with well-watered crops. By
contrast, Robertson et at. (1980) concluded that root length density in
groundnut was not affected by differential water management. Total root
length in a 150 cm profile was not significantly different (range of 68-
(a) (b)
Root length density (cm/cm3)

0 0.5 1.0 1.5 0 4 .8 1.21.62.024
15
20 30
E E
.3- 45
.3- .r:
.r: 40 i5. 60
i5. Q)
Q) "0 75
"0
·0
90
o WET
·0 60 CJ)
CJ)
105
80 120
15
20 30
E E
.3- .3- 45
.r: .r: 60
i5. 40 i5.
"0
Q) Q)
"0 75
~ DRY
·0 ·0 90
CJ) 60 CJ)
105
80 120
Figure 9.3 Root length density distribution for groundnut under well-watered and
water-stressed regimes at: (a) day 68; (b) day 55. (After Pandey et al., 1984b;
Devries et al., 1989a.)
108 cm/150 cm) between irrigation treatments. It is possible that the

relatively high rainfall (330 mm) which occurred during this experiment did
not cause equivalent crop water deficits, and hence root responses, to that
observed by Pandey et al. (1984b) and Devries et al. (1989a).
The growth rate of roots is markedly affected by soil water content
through its effect on soil strength and soil water potential (Klepper, 1987).
Groundnut tap root elongation decreased from 60 mm/day to less than
10 mm/day as soil strength, as measured by penetrometer resistance,
increased from 0 to 5 MPa (Taylor and Ratliff, 1969). Controlled environ-
ment studies indicate that root elongation in groundnut appears to be less
severely affected by compact soil than is cotton (Taylor and Ratliff, 1969),
pea (Gerard et al., 1972) and pigeon pea (Kirkegaard et al., 1992). Field
evidence also indicates that groundnut roots had superior root penetration
and development through a compacted soil zone when compared with
soybean and pigeon pea (Devries et al., 1989a).
Plant population can influence the rate of soil water extraction and final
rooting depth in groundnut. Nageswara Rao et al. (1989b) found that total
length and weight of the root system decreased with wider rows and
reduced plant density, although significant amounts of root biomass were
still invested in the inter-row spaces of the sparse plantings (12 plants/m2 ).
Soil water extraction from depth commenced sooner; it was more rapid in
the denser stands and was associated with greater root production at
deeper layers in the soil (Simmonds and Williams, 1989). Similar ground-
nut plant population effects on the pattern of water extraction and use in
response to crop water deficits imposed during early crop growth have
been reported elsewhere (Bhan and Misra, 1970; Ishag et al., 1985; Wright
and Bell, 1992b).
9.4.2 Shoot growth
(a) Vegetative growth

Soil water deficits reduce leaf and stem growth through effects on plant
water status, photosynthesis and leaf expansion (sections 9.2 and 9.3).
Boote et al. (1982) and Boote and Ketring (1990) have extensively
reviewed the literature on this aspect, and give more detailed descriptions.
In groundnut, reproductive growth consists of three distinct stages:
production of flowers, development of pegs that carry the ovary below
ground, and the subsequent formation and filling of pods.
(b) Flowering
Water deficits during flowering can result in a decrease in flower number
and a delay in time to flower. The timing and severity of stress largely
influence these effects (Boote et al., 1982). For instance, Chapman (1989)
showed that the initiation of flowers ceased within five days of applying an
early stress, at which time about 65% of extractable soil water had been
used. Since only 15-20% of flowers result in pods that contribute to yield
(Smith, 1954), reductions in flower numbers arising from water deficits do
not directly influence pod yield (Nageswara Rao et al., 1988). Also,
groundnut can compensate for reduced flower numbers resulting from
water deficits by producing a flush of flowers once the stress has been
relieved (Pallas et al., 1979; Nageswara Rao et al., 1988; Harris et al.,
1988).
(c) Pegging and podset

Soil water deficits during pegging and podset decrease yield primarily by
reducing pod number rather than kernel weight per pod (Skelton and
Shear, 1971; Ono et al., 1974; Boote et al., 1976; Pallas et al., 1979). Harris
et al. (1988) wisely point out that this is true only when sufficient water is
available later in the season to allow adequate production of assimilates.
Numerous experiments reporting the influence of mid-season drought yield
have been conducted, with general pod yield reductions in the range of
15-30% of well-watered controls (Pallas et al., 1979; Stansell and Pallas,
1985; Nageswara Rao et ai., 1985; Chapman, 1989; Wright et al., 1991).
A reduction in soil water content can have a dual effect on peg and pod
development owing to the subterranean fruiting habit of groundnut. On
the one hand, root zone water content can directly affect plant water
status, photosynthesis and hence assimilate supply to developing pegs and
pods. On the other hand, water content in the pegging and podding zones
(soil surface to a depth of 5 cm) can affect reproductive growth indepen-
dently of root zone moisture content. For instance, peg penetration
and conversion into pods, as well as pod calcium and water uptake, are
influenced by pod zone water content (Boote and Ketring, 1990). Because
the effects of soil water content on reproductive development are not
always separable, much of the research on this topic refers to soil water
status without reference to root or podding zones (Boote et al., 1982).
Considering the direct effect of soil water content first, Chapman (1989)
has shown that peg initiation and peg elongation in four groundnut culti-
vars ceased when about 80% of the extractable soil water had been
exhausted. Figure 9.4 shows the sensitivity of peg initiation and elongation
to soil water deficits applied during early reproductive development (46-67
days after planting, cv. Robut 33-1). Interestingly, pegs initiated prior to or
during the drought period had the capacity to renew elongation upon
rewatering. This attribute occurred in all four cultivars studied and appears
to be an important adaptation in intermittent drought situations. Stirling et
al. (1989b) studied the physiological mechanisms controlling the partition-
ing of assimilate to pegs and pods using labelled 14C. It was found that the
rate at which new assimilate was allocated to pegs declined before there
was a detectable fall in the turgor potential of pegs. This finding implies
that the primary mechanism restricting elongation of pegs during the
drought was indirect, and was probably mediated by plant hormones.
Cultivars differ substantially in their pegging and pod set response to soil
water deficits. Harris et ai. (1988) showed that the cultivar TMV-2 was able
to maintain a peg production efficiency (ratio of pod number to peg
number) of about 0.8 irrespective of a drought applied during the early
reproductive phase (17-72 days after planting). In contrast, three other
cultivars (Kadiri-3, NCAC 17090 and EC 76446 (292)) had an efficiency of
only 0.15 during the drought. Upon relief of the stress (73 days after
planting) all cultivars increased their efficiency to 0.8; however, Kadiri-3
took longer to reach this level owing to a ,renewed flush of flowers, pegs
and pods. These differing reproductive patterns in response to drought
provide useful information on the physiological basis of adaptation to
drought among cultivars. For instance, the yield strategy of TMV-2 was
to produce and fill pods at a moderate but constant rate, irrespective of
changes in drought intensity. In contrast, the drought-sensitive character
of Kadiri-3 resulted in a late flush of pods following rewatering which could
250
(0)
200
C
til
:9: 150 0
(j; /",--
0 .../
.0 0-0----·
E 100
:::J
0.......... /
' /
-.-.- .•
C
01 /
CD
Il. , /
50
0
50 60 70 80 90
Days after sowing
(a)
100 100
oA .8 oA e8
80
Ib) Vc
oE
'0
eF 80
Ie)
Vc '0
f'.G .6:H
oE A1-1
E E
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f
.r. .r.
0, 0,
c c
~
l!! l!! 40
01
40 01
CD CD
Il. Il.
20 20
0 0
50 60 70 80 90 50 60 70 80 90
Days after sowing Days after sowing
(b) (e)
Figure 9.4 (a) Production of pegs (cv. Robut 33-1) in response to non-limiting
water (0-0), water stress between 46 to 67 days after planting (.-.) and 61 to 78
days after planting (0-0). Elongation of pegs under (b) well-watered conditions
and (c) when water was withheld from 46 to 67 days after planting. Each symbol
represents a different age class, i.e. pegs that appeared on the same day. (After
Chapman, 1989.)
not mature prior to harvest. Similar cultivar responses have been reported
elsewhere (Wright et al., 1991).
We now consider the indirect effect of podding zone water content on
reproductive development. The situation where a dry surface soil overlays
a moist subsoil is particularly common in ground nut producing areas where
both intermittent and protracted droughts occur. Even where irrigation is
employed on sandy soils, adequate water may be applied to supply the root
zone for a period of a week or so but the surface podding zone may dry out
rapidly in only a few days. Very little research has been conducted to
understand the effects of dry podding zone soil on peg penetration and
subsequent development into pods. It seems· that peg and pod develop*
ment under these conditions may be influenced by podding zone water
content directly, by lack of calcium (Ca) uptake in developing pods in dry
soil, and by greater soil strength associated with dry soil.
A few investigators have used techniques to separate the rooting and
podding zones in order to study the effects of pod and root zone water
content on reproductive development (Bennett et al. (1990) give a review
of these techniques). Underwood et al. (1971) showed that, although pod
development was unaffected in podding zone soil maintained at higher
than -1.5 MPa, less than 20% of pods developed in air*dry soil compared
with well-watered control plants. Skelton and Shear (1971) found that a
dry podding zone significantly reduced the number of pegs which devel-
oped into pods, as well as increasing the number of dead pegs. Wright
(1989) demonstrated the existence of cultivar differences in peg and pod
development in response to poJding zone water content. Where the root
zone was kept moist and the podding zone was air-dry, seed yield of cv.
Robut 33-1 was the same as a treatment whose podding zone was kept at
field capacity, while seed yield in cvv. McCubbin and Gajah was reduced
by 43% and 35% respectively. A decrease in the number of pegs that
developed into fully grown pods along with a reduction in the number of
seeds per pod were responsible for the yield reductions. These results
suggest that selection for cultivars capable of maintaining normal repro-
ductive development in dry podding zone soil may be possible. Bennett et
al. (1990) found that a dry podding zone reduced the percentage of cv.
Florunner pegs which developed into full pods from 61 % to 48%, and also
reduced seed yield by 27% relative to the well-watered control plants.
It is well documented that low podding zone water content can decrease
Ca uptake and thus induce Ca deficiency in groundnuts (Hallock and
Allison, 1980; Rajendrudu and Williams, 1987). The interaction between
soil water deficit and Ca requirement for developing pods occurs as a result
of the subterranean nature of groundnut fruits. In aerial fruiting plants, Ca
is absorbed by roots and transported via the xylem vessels to the fruit.
After being conducted upwards and absorbed in the shoot, it then remains
highly immobile (Hanson, 1984). In groundnut, early studies showed that
Ca cannot move from one side of the plant to the other, or be translocated
from the root to the developing pod (Bledsoe et at., 1949; Brady, 1947).
The phloem is unable to conduct sufficient Ca to the developing carpo-
phore, although it does conduct all other substances necessary for pod
growth (Wiersum, 1951). After carpophore enters the soil, the developing
pod has no functional evaporative surface, and therefore no direct access
to root-absorbed, xylem-transported Ca (Skelton and Shear, 1971;
Wiersum, 1951; Beringer and Taha, 1976). The Ca requirement for devel-
oping pods is therefore mainly derived from absorption (through the pods)
direct from the soil solution.
Considering the requirement for direct soil uptake of Ca to developing
pods, it is difficult to explain how groundnut pods develop at all under very
dry pod zone conditions. Bennett et at. (1990) suggest that most of the
available evidence supports the hypothesis that carpophores enter a dry
podding zone and lose water by transpiration from peg or developing pod.
Under these conditions Ca can move in the transpiration stream to the
developing fruit. Underwood et at. (1971) suggests that this pathway may
provide sufficient water (and presumably Ca) for normal pod development
down to a soil water potential of -1.5 MPa, but not in air-dry soil. The
physiological mechanisms underlying Ca uptake and pod development in
dry soils is not well understood. The fact that cultivar differences in pod
development and seed calcium concentration in air dry pod zone soil exist
in groundnut (Wright, 1989) means that comparative physiological studies
to elucidate such mechanisms should be possible.
Calcium, usually applied as gypsum, has been widely shown to increase
yield of groundnut under drought conditions occurring during early pod set
(Hallock and Allison, 1980; Cox et at., 1976; Rajendrudu and Williams,
1987; Radder and Biradar, 1973; Balasubramanian and Yayock, 1981).
Rajendrudu and Williams (1987) suggested that gypsum application, which
resulted in more rapid establishment of pod numbers, provided an effec-
tive drought escape mechanism by getting more pods past the critical pod
set sage. Under well-watered conditions, this effect can provide greater
pod synchrony and even more maturity.
The inability to relate groundnut yield response to soil test Ca levels
across a wide range of soil water availabilities (Daughtry and Cox, 1974) is
no doubt related to reduced Ca availability and uptake in dry podding
zones. Large-seeded virginia cultivars have frequently been shown to be
more sensitive to drought-induced Ca deficiency than smaller seeded types
(Stansell et at. 1976; Slack and Morrill, 1972; Beringer and Taha, 1976).
The sensitivity of the large-seeded cultivars to Ca supply was suggested to
be related to the smaller surface-to-volume ratios of the large pod when
compared with a small pod (Boote et at., 1982). Kvien et at. (1988) later
confirmed that this was the case by studying the influence of several pod
characteristics on Ca accumulation and concentration in eight groundnut
genotypes. The ability of a cultivar to translocate Ca from shells to
developing seeds also appears to be important in determining cultivar
sensitivity to soil Ca levels. Beringer and Taha (1976) found that, although
the large-seeded cultivar, Makulu Red, absorbed more Ca than the smaller
seeded Natal Common, it was less effective in translocating Ca from shells
to seeds. Similarly, Kvien et al. (1988) concluded that thin, light shells trap
less Ca than thick, dense shells, and hence promote higher Ca concen-
trations in the seed. Recent investigations by Webb and Hansen (1989)
have also shown that carpophores and pods produce unicellular structures
resembling root hairs up to 0.75 mm long, which reach very high densities.
These structures, which were evident at the R5-6 growth stage (Boote,
1982) and degenerated by maturity, are considered to be an adaptation for
calcium and water uptake. Wright (1989) also found that cv. Robut 33-1
had more pod hairs and higher seed Ca concentrations compared with two
cultivars whose pods were grown in air-dry soil. It was hypothesized that
these hairs may have increased the effective pod absorptive surface area
and facilitated Ca uptake under drought conditions.
Surface soil physical condition is of considerable importance to peg and
pod development in groundnut, considering that the peg must penetrate
the soil surface to a depth of 2-7 cm to allow normal pod development
(Smith, 1954). It is surprising that there is only one report in the literature
pertaining to this topic. Underwood et al. (1971) showed that, with soil
water at field capacity, the ability of pegs to penetrate depends on the bulk
density of the soil. Figure 9.5 shows that the number and total weight of
pods per plant decreased asymptotically as penetration resistance (and
bulk density) increased. They also showed that most pegs which penetrated
1.0-1.5 cm developed pods, although development was slower when pods
were near the soil surface. This solitary set of data clearly establishes that
soil physical factors have a marked influence on peg penetration and
subsequent pod development.
Wright et al. (1992) studied the influence of surface soil strength on
reproductive development in groundnut. Pegs were allowed to enter small
cores which had been packed to different bulk densities. The depth of peg
entry and pod growth (dry weight) were measured in two cultivars (Robut
33-1 and McCubbin) 21 days after pegs were introduced into the cores.
Figure 9.6 shows the relationship between depth of peg penetration and
soil strength, as measured by penetrometer resistance. It was clear that
depth of peg penetration (and subsequent pod growth) declined linearly as
penetrometer resistance increased from 0.1 to about 2.0 MPa, with pegs
ceasing to penetrate and develop further at resistances greater than
2.0 MPa. There was little evidence of cultivar differences in peg response
to soil strength; however, further experiments assessing cultivar variation
in the 0-2.0 MPa region need to be conducted with greater precision. The
peg response to soil strength agrees well with the data of Underwood et al.
(1971), which showed the number of developed pods, and pod yield per
plant, ceased at penetrometer resistance> 2.0 MPa.
Figure 9.6 also shows how pegs appear to be considerably more sensitive
280 (a) 280

(b)
!II
240 240
C>
0)
a. 200 200
ala. E
.!!!
160
0
Qj
>
~
!II
160
8.'0
0)
-0
120 120
'0
:D \.
.0 80 J2
E 0)
80
>=
z" • •
• •••
40 40
0 • 0
2 3 4 5 6 2 3 4 5 6
Penetrometer force (MPa) Penetrometer force (MPa)
Figure 9.5 (a) Pod yield per plant and (b) number of pegs that developed to
diameters greater than 3 mm, as a function of penetrometer resistance in the
surface 1.5 cm of soil. (Reproduced with permission, Underwood et al., 1971.)
80
• Root elongation
70 0 Robut 33-1 3.0
i McCubbin
60
2.5 :c
..,
0
E
.s 50 \\ , 2.0
~
(1)
0"
....... . "a
~ 00
E co
40
.
0)
C>
o '
,,~ 1.5 o·
0)
a. 30 "Dl
'0
.<:;
""- "- 1.0
CD
0.
0)
0
20
io
""- "- "- • "3
~
.•
::T
"- 0.5 .;:!.
10 "-
"- -.
......
oi
0
or
o i
0
0 2 3 4 5 6 7 8 9 10 12
Penetrometer resistance (MPa)
Figure 9.6 Depth of peg entry as a function of penetrometer resistance in the

surface 1.5 cm of soil for two cultivars measured during a 21-day pegging period
(Wright and So, unpublished data). The broken line shows the effect of penetrom·
eter resistance on root elongation (after Taylor and Ratliff, 1969).
to soil compaction than do roots, as Taylor and Ratliff (1969) showed
groundnut roots ceased growth at a penetrometer resistance of >5.0 MPa.
The results of this experiment have demonstrated that peg entry, pod
growth and development are extremely sensitive to soil hardness, which
commonly increases with declining surface soil water content (Sivakumar
and Sarma, 1986; So and Woodhead, 1987). It therefore follows that high
soil strengths during the pegging and pod set phase must severely inhibit
reproductive efficiency and reduce harvest index and pod yield in
groundnut.
(d) Pod filling

Water deficits during pod fill generally reduce pod and kernel weight
(Pallas et al., 1977, 1979; Pandey et at., 1984a; Lenka and Misra, 1973;
Nageswara Rao et al., 1985; Wright et al., 1991). The timing of water
deficit in the pod filling stage largely influences pod growth, pod size and
maturation. There are numerous reports available on the effect of drought
during podfilling but the findings vary enormously (as reviewed by Boote et
at., Boote and Ketring, 1990). This is not surprising, considering that some
work has investigated the effect of water deficits within each growth phase
separately, with water being non-limiting during earlier phases (e.g.
Nageswara Rao et al., 1985), while other studies have considered pro-
tracted water deficits throughout the season (Mathews et at., 1988a). There
are therefore many possible interactions between plant water status and
stage of growth, which influence the establishment of pod numbers and the
availability of assimilate to fill those pods.
Late-season drought in longer-season maturity types has been shown to
reduce pod yields more severely than in shorter-season types, largely
through reductions in pod numbers and seed size relative to well-watered
controls (Boote et at., 1976; Pallas et al., 1979; Wright et al., 1991).
Shorter-season types are able to initiate and develop pods earlier and
thereby escape severe soil water deficits late in the season.
Water deficits occurring during the pod formation stage can delay the
establishment of a full pod load, as discussed in the previous section. Upon
relief of this stress, groundnut has often been observed to initiate a flush of
flowers, peg and pods which can delay harvest maturity by 10 or more days
(Stansell and Pallas, 1985; Boote et at., 1976; Wright et al., 1991; Shorter
and Simpson, 1987). In such a situation, two or more flushes of pod
formation can occur, and farmers using mechanical harvesters may need to
make decisions on which flush of pods is the most suitable to harvest. In
many environments full yield compensation through delayed harvesting
may be difficult, as overmature pods may remain in the soil, and in
subtropical environments low temperatures late in the season may reduce
pod growth rates and hence quality (Bell et al., 1992). Cultivars which have
more synchronous pegging and podding patterns, despite the occurrence
of soil water deficits, can overcome this problem (Harris et al., 1988;
Chapman, 1989; Wright et al., 1991).
9.4.3 Effects of water deficits on partitioning and harvest index

Harvest index (H), defined as the proportion of pod to total biomass, can
vary enormously depending on the timing and severity of water deficit
relative to pod set (Ong, 1986). Duncan et al. (1978) suggested that H can
be considered as a function of the crop growth rate (CGR), partitioning of
assimilate to pods (P), and the effective duration of podfilling phase (t).
Mathews et al. (1988a) showed that H can be expressed as the mean value
over the season of p weighted by CGR, described by the expression:
H = (p CGR t)
(1)
W
where W is total biomass, adjusted for energy content in the pods (Duncan
et al., 1978), at maturity. This simple analysis allows the investigation of
specific variables influencing treatment and cultivar differences in H in
response to water supply. The above framework has been used by a
number of workers to analyse cultivar and treatment differences in ground-
nut pod yield performance under a range of soil water regimes. For
instance, Duncan et al. (1978) concluded that cultivar differences in p,
rather than in CGR and t, were largely responsible for pod yield improve-
ment of recently developed cultivars in the USA under well-watered
conditions. Mathews et al. (1988a) and Harris et al. (1988) found that
cultivar differences in pod yield during a terminal drought were due to
variation in H, which was associated with differences in p and t. Similarly,
Wright et al. (1991) showed that CGR was similar among four cultivars;
however, differences in p and t were responsible for H and pod yield
variation in a terminal drought situation. The positive pod yield response
to drought during the pre flowering phase, compared with a fully irrigated
treatment, observed by Nageswara Rao et al. (1985, 1988) could largely be
explained by greater synchrony of pod set which allowed a longer period of
podfilling (i.e. greater t). There were only minor differences in p and CGR
among irrigation treatments.
During podfilling the size, number and strength of reproductive sinks,
relative to other sinks in the plant, determine the final degree of partition-
ing to pods. It has been shown in many determinate crops that pre-anthesis
biomass can be remobilized to reproductive sinks from other parts of the
plant, particularly in response to water stress (e.g. Constable and Hearn
1978; Bidingeretal., 1977; Blumetal., 1983; Wrightetal., 1983a). Ludlow
and Muchow (1988) suggest this trait can improve yield stability by acting
as a buffer against the effects of water deficits on current assimilation.
Duncan et al. (1978) and Ong (1986) have suggested that no translocation
of vegetative assimilate to pods occurs in groundnut, as partitioning ratios
(PR, ratio of pod growth rate to crop growth rate) were always less than
100% (Table 9.1). In contrast, Bell (1986) recorded PRs well in excess of
100% in irrigated groundnuts (cv. Early Bunch) in a tropical environment,
which suggested that substantial remobilization of assimilate can occur.
Other recent data (Chapman, 1989; Wright et al., 1991) has shown PRs in
excess of 100% can occur, and also that cultivar differences in PR were
apparent in groundnut subjected to water deficits during the podfilling
phase (Table 9.1). Based on this data, cultivar variation in the degree of
remobilization during podfilling may exist, and more research into its role
in improving H and pod yield in groundnut under drought conditions
seems warranted.
9.5 FACTORS AFFECTING EVAPOTRANSPIRATION
Evapotranspiration (E1), or crop water use, refers to the combined pro-

cesses of evaporation and transpiration which account for the consumptive
water loss during crop growth. While transpiration (1) is responsible for
water used by the plant, soil evaporation (Es) represents water loss from
the soil surface that does not pass directly through the plant. While T may
be readily measured in container studies in which Es can be minimized, it is
much more difficult to measure, calculate or estimate soil evaporation in
field studies (Turner, 1986a). Most field data that have been reported for
groundnut simply combine T and Es. This has made generalizations about
crop production as related to ET, rather than T, a very difficult task
(Ritchie, 1983). Boote et al. (1982) have discussed experimental techniques
available for estimating groundnut ET in the field.
ET is influenced by many factors, which can be broadly categorized into
the effects of canopy development and soil water deficits. These are briefly
discussed below and have been exhaustively reviewed by Boote et al.
(1982) and Boote and Ketring (1990).
9.5.1 Effect of canopy development on ET

Stansell et al. (1976) showed that daily ET in groundnut increased from
2 mm/day at 20 days after planting (DAP) to a maximum of 6 mm/day at
between 70 and 95 DAP, depending on cultivar. ET then declined gradu-
ally to about 3 mm/day at maturity, in response to plant senescence and
decreasing evaporative demand. Similar seasonal ET responses have been
observed elsewhere (Dancette and Forest, 1986; Ishag et ai, 1985). Figure
9.7 shows a typical seasonal pattern of daily ET for a virginia type (cv.
Ashford) grown under well-watered and water-limited conditions in the
Sudan, Africa (Ishag et al., 1985). Higher maximum daily ET rates of
7-10 mm/day have been recorded under conditions of high evaporative
demand (Mantell and Goldin, 1964; Grosz, 1986).
Factors affecting evapotranspiration 303
The seasonal pattern of ET in groundnut is closely related to the pattern
of canopy, and hence LAI, development (Ishag et ai., 1985: Dancette and
Forest, 1986; Wright et ai., 1991). When crop canopy is establishing, a
large proportion of ET can be lost through soil evaporation (Es) (Cooper et
ai., 1983). Ritchie and Burnett (1971) found that in the early stages of crop
TABLE 9.1 Partitioning ratios (PRJ in groundnut cultivars, as influenced by water

supply
Cultivar Remarks PR
(%) Reference
Dixie Runner Irrigated 40.5 Duncan et al.

(1978)
Early Runner 757 "
Florunner 84.7
Early Bunch 97.8
Span cross 66.2
Early Bunch Irrigated, planted 7/12 91.3 Bell (1986)

29/12 85.7
18/1 128.7
8/2 125.0
1/3 143.5
Virginia Bunch Terminal drought 82.0 Wright et al.

(1991)
UF78114-1 110.0 "
McCubbin 88.0
Red Spanish 88.0
Virginia Bunch Irrigated 99.0 Chapman (1989)

McCubbin 107.0
UF78114-1 107.0
Robut 33-1 98.0
Virginia Bunch Early stress 330

McCubbin 25.0
UF78114-1 50.0
Virginia Bunch Late stress 102.0

McCubbin 122.0
Robut 33-1 118.0
FRs were calculated as the ratio of pod growth rate (adjusted for oil content) to crop growth
rate x 100.
growth, until an LAI of 1.0 is reached, Es can account for well over 50% of
the total ET. Simmonds and Williams (1989), for instance, found that
when groundnut canopy coverage was small, about 16 mm was lost by soil
evaporation during one week following an irrigation, with almost half of
this loss occurring in the first two days. The rate of Es is dependent on
surface soil wetness, and the degree of shade provided by the canopy.
When the surface soil is very wet, the rate of evaporation is governed by
the supply of energy, and upon further drying Es is limited by soil hydraulic
conductivity (Philip, 1957).
To obtain greater understanding of the effect of management or cultivar
on crop water use, it is necessary to split ET into its components - namely
T and Es. The direct measurement of Es using small trays filled with fully
drained soil and sunk in between rows has been attempted in groundnut
(Simmonds and Williams, 1989). Trays were removed daily and weighed,
with the difference in weight being assumed to be water lost by evapor-
ation. The technique is unreliable for estimating water loss after the first
few days after wetting (ODA, 1987). Ritchie (1972) and Tanner and Jury
7
(a)
6
2
>:
III 01"·
~
S 6
I- (b)
w
5
3
2
0<
5 15 25 35 45 55 65 75 85 95 105 U5
Days after sowing
Figure 9.7 Mean daily evapotranspiration as a function of time under (a) well-
watered and (b) water-limited conditions, in (.) dense and (0) sparse stands of
groundnut. (After Ishag et at., 1986.)
Yield responses to evapotranspiration/transpiration 305
(1976) developed empirical models to describe the temporal change in
water loss from an uncropped soil and the effect of crop canopy develop-
ment on radiant energy interception. These models are predictive in
nature, and have been successfully used in larger crop models (e.g. Boote
et al., 1986). Cooper et al. (1983) extended this approach, and described a
technique whereby standard field measurements of ET, evaporation from
uncropped soil and canopy cover (via canopy light interception, or LAI
and canopy extinction coefficient measurements) allow reliable estimates
of Es and T. Wright et al. (1994) used this technique to estimate T and E in
mini-lysimeters installed in groundnut canopies in the field. In a terminal
drought stress treatment, Es was 45 mm out of a total ET of 185 mm, when
averaged over four cultivars.
9.5.2 Effect of soil water deficits on ET

ET will continue at an energy limited rate until water content of a large
part of the root zone drops below a critical level, when stomatal closure
restricts water loss. There is little published information on the critical
level of available water content below which ET declines in groundnut.
Wormer and Ochs (1959) found that T of groundnut remained constant
until two thirds of the available soil water had been depleted, after which it
decreased to the wilting point. In a range of crops and locations, Ritchie et
ai. (1976) found that the ratio of ET to potential evaporation from freely
evaporating surface remained relatively constant if extractable soil water
was greater than 25% , then decreased linearly between 25% and 0%. The
generality of this response across a wide range of environments has been
questioned. Following the theoretical analyses by Jordan and Miller (1980)
it was shown that the decline in ET is reached at higher soil water contents
under high evaporative demand conditions because steeper gradients of
soil water potential are required to sustain flow. Denmead and Shaw
(1962) demonstrated this effect with corn plants grown in containers in the
field under differing meteorological conditions. The implications of these
findings are that the magnitude of the total extractable soil water varies,
depending on the evaporative conditions experienced by the crop. It also
depends on rooting depth, root length and density, and root resistance
(Sheriff and Muchow, 1984).
9.6 YIELD RESPONSES TO EVAPOTRANSPIRATION AND

TRANSPIRATION
Based on a review of the available literature, Boote et ai. (1982) and Boote
and Ketring (1990) concluded that about 600 mm of water is required for
optimal pod yield performance. The presented relationships between pod
yield and ET showed significant variability, which they attributed to factors
other than water use. An example of the type of variability observed is
illustrated in Figure 9.8(a), which compares the relationship between pod
yield and ET for data from Georgia, USA and Hyderabad, India. While
pod yield increased linearly from approximately 1.5 t/ha to 6 tlha in both
regions, the amount of ET used differed substantially, ranging from 200 to
600 mm for the USA data, compared with 450-800 mm for the Indian data.
Hanks (1983) and Kanemasu (1983) discussed some of the factors influenc-
ing the pod yield/ETrelationship in a range of crops and concluded that (a)
improvements in the correlation are observed when T rather than ET is
used; (b) total dry matter production (TDM) and ET are more closely
correlated than pod yield and ET; and (c) pod yield/ET relationships can
vary from location to location, year to year and crop to crop. These factors
as they relate to groundnut yield/water use relationships are briefly dis-
cussed below.
9.6.1 Transpiration versus evapotranspiration

De Wit (1958) concluded there was a close relation of TDM to T across a
wide range of species and conditions. This response is not surprising: TDM
and photosynthesis are closely related, and photosynthesis and transpi-
ration are also related via the diffusional resistances (Tanner and Sinclair,
1983). In field studies, the problems of evaluating the amount of Tfrom the
generally measured ET (soil evaporation plus T) can introduce consider-
able error into T estimates, and hence the ET/TDM relationship.
Similarly, drainage below the root zone and/or runoff can cause consider-
able error in ET measurements. (Methods of estimating T from ET
measurements are discussed in section 9.5.)
9.6.2 TDM versus pod yield

Figures 9.8(a) and (b) illustrate how there was a considerably stronger
correlation between TDM and ET (,2 = 0.61) compared with pod yield and
ET (,2 = 0.38) for groundnut crops exposed to different timings of soil
water deficits (Nagweswara Rao et at., 1985). Indeed in a narrow range of
nearly the same total ET (500-600 mm), pod yield varied from 0.8 tlha to
over 5 t/ha, depending on the timing of water deficit. Stansell and Pallas
(1985) also showed that different pod yields occurred at similar total ET
values, as a result of timing of water deficits. Attempts have been made in
other crops to relate harvest index to the timing and severity of water stress
in order to improve the prediction of ET/pod yield relationships (Slabbers
et at., 1979; Stewart et at., 1977). Kanemasu (1983) warns that the ETlpod
yield relationships are not unique because of the complex interactions
between development, assimilate partitioning and environment, and con-
siders it is doubtful that an ET/pod yield relationship can be extended to
climatically diverse regions.
Yield responses to evapotranspiration/transpiration 307
6,0
(a) • •
• •• • •
5,0 • • •
• •
••
4,0 •
<? • •
~ ••
:Q 3,0
Q)
's;,
-0
o
:
a..
2.0
•
•
• •
•
1.0
•
•
0.0
o 100 200 300 400 500 600 700 800
ET (mm)
11.0
(b)
10.0 •
•
9,0
8,0
7,0
<?
~ 6,0
Q;
'iiiE 50
'
~ 4,0 •
(ij
•
~ 3,0
2.0
•
1.0
0.0
o 100 200 300 400 500 600 700 800
ET (mm)
Figure 9.8 Relationship between (a) pod dry matter and evapotranspiration (ET)
for data derived from (-) Stansell et al. (1976) and (.) Nageswara Rao et al. (1985);
and (b) total dry matter and evapotranspiration for data derived from Nageswara
Rao et al. (1985).
9.6.3 Environmental effects
Much of the variation in the ETlpod yield relationship between the USA and
Indian crops shown in Figure 9.8(a) can be explained by site differences in
evaporative demand. Ong et al. (1987) showed that in groundnut T
increased, and TDM increased linearly as vapour pressure deficit increased
from 1 to 2.0 kPa. Differences in evaporative demand from year to year have
been shown to change the ETlpod yield relationship in other crops (Hanks,
1983). Other possible reasons for variations in the relationship between ET
and pod yield include diseases, insects, nematodes, nutritional stress, soil
water capacity and rooting depth. Boote and Ketring (1990) suggest that a
modelling approach capable of integrating these interacting effects may
enable better prediction of pod yield response to water application.
9.6.4 Water use efficiency and transpiration efficiency

Water use efficiency can be looked at from many perspectives, and care is
needed in defining its use. Water use efficiency (WUE) is generally defined
as dry matter production (total or pod) per unit of ET, while transpiration
efficiency (TE) is dry matter production per unit of crop transpiration. The
distinction between these terms is important in the context of genetic and
agronomic solutions to the problem of drought in groundnut. For instance,
treatments that suppress soil evaporation can improve WUE but need not
improve TE (Tanner and Sinclair, 1983). Several models have been suc-
cessfully employed to predict dry matter production from ET and T in a
range of crops (de Wit, 1958; Arkley, 1963; Bierhuizen and Slatyer, 1965;
Tanner and Sinclair, 1983). Such models allow opportunities for improving
the efficient use of water to become more clearly defined. All of the models
are largely derived from the analysis of de Wit (1958) which proposed that,
in closed canopies:
TDM m
(2)
T Eo
where Eo is the potential evaporation from pan data or the Penman
equation, and m is a constant governed mainly by species. The later models
used vapour pressure deficit rather than Eo to improve prediction across
contrasting environments. Tanner and Sinclair (1983) concluded that WUE
can mainly be improved by modifying the vapour pressure environment in
which plants grow by, for instance, applying mist irrigation or planting
crops earlier or later to coincide with lower vapour pressure deficits.
Similarly, management options such as plant population which modify
evaporation loss relative to transpiration can influence WUE, with little
effect on TE.
It has been recognized for many years that the internal method of
utilizing carbon can markedly influence TE of plants with C4 plants having
Ameliorating the effect of water deficits on yield 309
about twice the TE of C 3 plants (Tanner and Sinclair, 1983). The constant
m in the above equation is used to account for such variations. Reviews of
the literature often concluded that the exploitable variations in TE among
cultivars within a species is small and the potential for improvement by
breeding is limited (Fischer and Turner, 1978; Fischer, 1981; Tanner and
Sinclair, 1983). Recent evidence in groundnut (and many other species)
has overturned this conclusion, with Hubick et at. (1986) reporting that the
variation in TE of up to 60% exists in a range of groundnut cultivars grown
in large pots in the glasshouse. Large cultivar differences in TE have
subsequently been demonstrated in the field under well-watered (Wright et
at., 1988a) and water-limited conditions (Wright et at., 1994), using mini-
lysimeters installed in small canopies. Table 9.2 shows the extent of this
variation, with Tifton-8 having the highest TE range at 3.1-3.7 g/kg,
compared with the lowest values of 1.8-2.0 g/kg for Chico. Interestingly,
variation in TE among cultivars was largely due to differences in biomass
rather than to differences in water use. This result suggests that photosyn-
thesic capacity, rather than leaf stomatal conductance, dominates the TE
response in groundnut cultivars. Similar groundnut cultivar differences in
TE have been reported in the field by Mathews et at. (1988a). In their
study, cv. Kadiri-3 had the highest (2.17 g/kg) and cv. EC 76446(292) the
lowest TE (1.71 g/kg). It is clear that considerable scope exists to improve
TE and ultimately pod yield under water-limited conditions by selection
for this trait in breeding programmes.
9.7 AMELIORATING THE EFFECT OF WATER DEFICITS ON

YIELD
About 80% of groundnut production occurs in seasonally rainfed areas of

the semi-arid tropics, where climate is characterized by low and erratic
rainfall. Drought is recognized as one of the major constraints for ground-
nut productivity in these regions (Gibbons, 1980). Although effects of
supplementary irrigation on yields are significant, this option is only feas-
ible on a small proportion of the total arable area. Therefore other
methods of improving or stabilizing groundnut yields in drought-prone
environments are necessary. Genetic and management options to alleviate
effects of water deficits on groundnuts are considered in this section.
9.7.1 Genotypic improvement

Despite the importance of drought as one of the major constraints to
groundnut production, the research effort into genetic improvement of
drought tolerance in groundnut has been limited. This situation has largely
arisen because of a lack of simple screening technologies to assess the
drought tolerance of genotypes, and a lack of access to a large groundnut
TABLE 9.2 Biomass (inc!. roots), water use and TE in groundnut cultivars under
well watered conditions (Wright et aI., 1988a) and two levels of water limited
conditions (Wright et aI., 1992b)
Study Cultivar Biomass Water use TE

(g) (kg) (g/kg)
Well-watered Tifton-8 63.1 17.0 3.71

VB-81 46.9 16.2 2.90
Robut 33-1 55.3 19.0 2.91
Shulamit 51.6 16.8 3.07
McCubbin 48.6 16.9 2.88
Cianjur 43.4 16.3 2.66
Rangkasibitung 41.6 16.9 2.46
Pidie 47.3 16.6 2.85
lsd P = 0.05 7.0 1.5 0.3
Water-limited (intermittent
stress) Tifton-8 37.5 12.2 3.07
Shulamit 35.7 12.8 2.79
McCubbin 36.3 13.4 2.71
Chico 20.5 11.4 1.80
(Terminal stress) Tifton-8 31.3 10.0 3.13
Shulamit 29.0 9.9 2.93
McCubbin 26.8 10.0 2.68
Chico 17.8 8.8 2.02
lsd P = 0.05 5.6 1.9 0.4
Measurements of the change in biomass and water use were between 40 and 90 DAP.
germplasm pool. Bidinger et al. (1982) and Garrity et al. (1982) argued that
genetic improvement in yields of crops can be brought about if attributes
that confer yield advantage under drought conditions can be identified and
used as tools in breeding programmes to enable identification of drought-
tolerant genotypes. Many traits (physiological and biochemical) have been
proposed for improving the yield performance of crops under drought
conditions (Seetharama et al. 1983; Turner, 1986a). There are, however,
few examples where the trait-based approach has been successfully
employed in large-scale drought resistance breeding programmes. Ludlow
and Muchow (1988) critically evaluated traits for improving crop yields in
water-limited environments.
(a) Screening for drought tolerance

Drought is a complex syndrome with three main components: its timing,
duration and intensity, which can all vary widely during a cropping cycle.
The extreme variability in these three components has made it difficult to
define plant attributes required for improved performance under all
drought situations. The success of a crop genotype in drought situations is a
syndrome of effects manifested by the genotypic attributes contributing to
the productivity. In addition, droughts interact with other problems,
enhancing or reducing their significance. For example, the involvement
and interaction of drought in calcium deficiency, induced seed abortion,
pod rots and aflatoxin contamination in groundnut are well recognized.
This complexity means that the plant breeder relies on an approach which
seeks to select genotypes solely on the basis of increased yield and stability
in drought-prone environments.
Systematic attempts to screen groundnut genotypes for drought toler-
ance have only been made at IRHO, Senegal (Gautreau, 1967, 1969, 1970;
Bocklee-Morvan, 1983), and at the International Crops Research Institute
for the Semi-Arid Tropics (ICRISAT) in India (ICRISAT, 1983). A
common approach has been used in both screening programmes, with yield
performance of genotypes as the major selection criterion. This approach is
not surprising since no simple screening traits for the identification of
drought-tolerant genotypes have been developed to date. The breeding
methodologies adopted by the researchers have varied substantially. For
example, Gautreau (1967) evaluated genotypes in Senegal, under rainfed
conditions in drought-prone environments. At ICRISAT, genotypes are
evaluated for drought tolerance based on their total dry matter and pod
yield performance under a range of water deficits created by a line-source
sprinkler (Hanks et ai., 1976) during critical growth stages.
Each approach has its limitations. In the Senegal programme, only
limited numbers of entries could be included in multi-Iocational experi-
ments, due to the high costs of maintaining these trials. Further, variability
in environmental conditions between sites, arising from variation in
amount and distribution of rainfall, confounds the intrinsic genotypic traits
contributing to superior drought tolerance. The ICRISAT approach allows
large numbers of genotypes to be screened but the line-source technique
has some limitations. For instance, screening has to be conducted in
seasons where interference from rainfall is minimized (i.e. during the dry
season). Also strong wind during irrigation can influence the systematic
nature of water deficits created, requiring complex statistical techniques
for data analysis (Murarisingh et ai., 1991).
However, the line-source system offers certain advantages, in that it
allows large numbers of genotypes to be evaluated at varying intensities of
drought in a given environment. The sprinkler irrigation technique simu-
lates rainfall of varying intensities which subsequently wets the soil to
different depths, a factor that is particularly important for groundnut with
its subterranean podding habit. The genotypes identified as being superior
yielding types under this technique do, however, have to be re-evaluated
under rainfed conditions in drought-prone environments.
TABLE 9.3 Groundnut genotypes identified as drought tolerant by various
researchers
Research Varieties Botanical Dura- Reference

programme group tion
IRHO, Senegal 55-437 Spanish 90 Gautreau (1967,1969,

1970,1978)
73-30 Spanish 95
Te-3 Spanish 90 Bocklee Morvan
(1987)
Ts 32-1 Spanish 90
KH-241-D Spanish 90
73-33 Virginia 110
USDA, ARS, USA Tifton-8 Virginia 140 USDA, 1984
ICRISAT ICG-1697 Spanish 110 ICRISAT,1986
ICGV-86707 Spanish 110 ICRISAT, 1990
ICGV-86635 Spanish 110
Table 9.3 summarizes the list of genotypes identified as being drought

tolerant by the various research programmes in different countries. The
genotypes were selected as being drought tolerant based on their yield
performance under imposed drought or low rainfall situations.
9.7.2 Selection of genotypes for drought-prone environments

Because yield is the most important trait of a genotype, farmers prefer to
plant those that have high yield potential in normal to good rainfall years,
but which are also capable of producing reasonable yields in the event of
drought (i.e. types that have low sensitivity to drought). Nageswara Rao
et al. (1989a) examined the relationship between genotypic sensitivity to
drought and yield potential (yields achieved under adequately irrigated
conditions) in 22 groundnut genotypes (Table 9.4) under a range of single
and multiple drought patterns which were simulated using a line-source
sprinkler technique (Hanks et al., 1976). The sensitivity of the crop to a
given drought pattern was examined by regressing the mean pod yield of
the 22 genotypes against the water deficit created across the line source
gradient within each drought pattern (Table 9.5). The slope (Sd) of this
regression represents the sensitivity of the crop to water deficits within
each drought pattern. The results suggested that the crop is sensitive to
drought under almost all drought patterns, as indicated by the negative Sd
term. The magnitude of sensitivity to drought also varied among the
drought patterns. The crop's sensitivity to drought was relatively less when
drought occurred early in the season (PI in Table 9.5). It seems that, for
some drought patterns, there is little variation among genotypes in the
sensitivity to drought (Sd) other than that associated with yield potential
(Yp), while for other drought patterns the association between Sd and Yp
is weak (column r2 in Table 9.5). For the early and mid-season droughts,
correlation between Sd and Yp was not as strong as it was for the droughts
which occurred during the pod-filling phase (Figure 9.9). Thus, breeding
for drought resistance without sacrificing yield potential may only be
feasible for some drought patterns (for example, early and mid-season
droughts); in other cases, drought resistance may require the sacrifice of
yield potential in good environments.
In areas where the probability of end-of-season drought is high, the best
strategy may be to select for earliness to enable escape from this type of
drought. The ability of a genotype to recover from mid-season drought
when water again becomes available also plays a dominant role in genotype
adaptation to a drought pattern where deficits are relieved by intermittent
rains (Harris et ai., 1988; Nageswara Rao et ai., 1988). In this type of
drought, selection in well-watered environments is unlikely to identify
genotypes with greater recovery responses. Although early water stress
reduces initial shoot growth and development, synchronous renewal of
vegetative and reproductive development is often observed when the
drought is relieved (Stirling et ai., 1989a,b). An increase in 14C trans-
location into stem apices and pegs has been observed when a crop was
TABLE 9.4 Timing and duration of droughts applied to create .the different
drought patterns (Pn) (from Nageswara Rao (et al.,J989a)
Drought patterns
DAS+ PI P2 P3 P4 P5 P6 P7 P8 P9 PlO P11 P12
1 U U U U U U U U U U U U
29 LS U U U LS U LS LS U LS U LS
57 LS LS U U LS U LS U LS U U U
66 U LS U U LS U LS U LS LS LS LS
72 U LS U U LS U LS U LS LS LS LS
82 U U U U U LS LS LS U LS LS LS
93 U U LS LS U LS LS LS U U U U
100 U U LS LS U LS LS LS U U LS LS
111 U U LS LS U LS LS U LS U LS LS
118 U U U LS U LS LS U LS U LS LS
das+ = days after sowing

U = uniform irrigation using sprinklers
LS = line-source irrigation to create eight intensities of water deficit
re-watered following a drought during the early reproductive phase.
However, it was interesting to note that there was no interrelationship
between 14C partitioning and plant water status, which meant that the
changes in 14C assimilate partitioning to stem apices and reproductive
structures preceded any detectable changes in bulk leaf turgor levels.
These results suggest that factors other than cell turgor such as cell wall
extensibility (Barlow, 1986) and plant growth substances (Davies et ai.,
1986; Turner, 1986b; ICRISAT, 1990) may mediate recovery responses of
developmental processes. Significant variation among groundnut geno-
types in ability to recover from mid-season drought has been observed by
Harris et ai. (1988) (Figure 9.10). The physiological factors responsible for
genotypic variation in recovery patterns is unclear at this stage.
Another approach in the selection and identification of drought-adapted
genotypes is to model the yield of genotypes in drought-prone environ-
ments using historical weather data and information on genotypic sensi-
tivity to a given drought pattern (Bailey, 1990). It is argued that future
research in genotype selection lies more in improved modelling of cultivar
performance in varying environments. This approach, however, requires
basic and detailed understanding of the effect of environmental factors and
their interactions on crop growth and yield in order to assess the potential
value of new cultivars in drought-prone environments.
Current research is assessing variability among groundnut germplasm for
putative traits related to drought resistance which may ultimately be used
TABLE 9.5 Regression coefficients of the linear relationship

between yield potential (Yp) and genotypic sensitivity to water
deficit (Sd) in 22 genotypes (from Nageswara Rao et aI.,
1989a)
Drought Regression coefficients r2

pattern
PI Yp = 1.56-0.007 Sd 0.15 NS
P2 Yp = 1.50-0.014 Sd 0.51 *
P3 Yp = 0.94--0.013 Sd 0.64**
P4 Yp = 0.95-0.013 Sd 0.87**
P5 Yp = 1.67-0.011 Sd 0.52*
P6 Yp = 0.003-0.013 Sd 0.91**
P7 Yp = 0.14--0.Q17Sd 0.91**
P8 Yp =-0.30-0.008 Sd 0.38 NS
P9 Yp = 1.61-O.016Sd 0.90**
PlO Yp = 1.57-0.015 Sd 0.86**
Pl1 Yp = 1.30-0.017 Sd 0.92**
P12 Yp = 0.15-0.015 Sd 0.82**
*, significant at 5% level
**, significant at 1% level
80
(a) 6 66
6 6 66 66
70 6 6 6
~ 6
6. 66. 6
6
"0 66. 6 6
Q5 66. 6.
.:;' 11 6 1111 11
"0
0 60 f1 t, t, 6 6 6
c.. ti 6 6 6.6
.£ 6. l' f1
(/)
(/) t, 6.6.
6.
0
...J
50 6 11
0~
f1 6
40
6.
30
100 200 300 400
Pod yield when irrigated (g/m2)
90
(b)
80 11
f
6 A
AI1 f.:.
6~6A66
"0
Q5 6 f.:.
.:;' 70 l:::. l:::.
6. 11
A
"0 6 6
0 l:::.
c.. 6 M 6~6 6 l:::.
.£ 60
(/) 11 l:::. 6
(/) l:::. 11
0 tJ.
...J
0~ tJ.
50 /:,
tJ.
40
300 400 500 600 700
Pod yield when irrigated (g/m2)
Figure 9.9 Relationship between % pod yield loss due to drought and pod yield
potential under irrigated conditions in 60 groundnut genotypes. Yield loss due to:
(a) midseason drought; (b) end-of-season drought. (After Nageswara Rao et al.,
1989a.)
50
Irrigated Robut 33-1
40
~ I I
~ 30
Q;
.0 / / TMV-2
E
:;J
..-- ,/ ." NCAC 17090
"0'" 20
/-- -- ......... /
/ :'='=:~ - -
0
0- f
I I EC 76446(292)
10 .. /
./
-.. '-"-':~':::':':/
/
/
.. ,.........:. ... .:..:..:..: ... ;;.-. .
0
70 80 90 100 110 120 130 140
Days after sowing
Figure 9.10 Number of pods developed over time by four groundnut genotypes
during, and following release from, an early season drought, ICRISA T Center,
post-rainy season, 1982-83. Standard errors are presented as vertical bars. (After
Harris et al., 1988.)
as selection criteria in breeding programmes. For instance, substantial

genetic variation exists for partitioning of dry matter to pods. (Mathews et
al., 1988a; Nageswara Rao et al., 1992), and efficient root systems with
capacity to penetrate deeper soil layers (Ketring et al., 1985; ICRISAT,
1990; Wright et al., 1991; Wright and Nageswara Rao 1993). As discussed
in section 9.6.4, recent evidence has also shown that large cultivar differ-
ences in transpiration efficiency (TE) exist in groundnut grown under
glasshouse and field conditions (Hubick et al., 1986, 1988; Wright et al.,
1988a, 1994). The difficulty in accurately measuring TE in glasshouse and
field situations means that it is virtually impossible to include such a trait in
large-scale breeding programmes. This difficulty may have been overcome
following the elegant work of Farquhar et al. (1982), who proposed on the
basis of theory that TE and isotopic discrimination against l3C during CO 2
assimilation ~ should be correlated in plants with the C3 photosynthetic
pathway. Subsequent glasshouse and field experiments have show that TE
and ~ are indeed well correlated in groundnut (Hubick et al., 1986, 1988;
Wright et al., 1988a, 1993a, 1994) and have raised the poss.ihility of using ~
as a rapid, non-destructive and relatively inexpensive technique for selec-
tion of TE in groundnut breeding programmes. Other related research has
shown that specific leaf area (SLA, cm2/g) is well correlated with TE (and
~) over a wide range of cultivars and environments (Nageswara Rao and
Wright, 1994). This observation has now raised the possibility of using
SLA as an even more rapid and inexpensive technique for selection of TE.
Continuing research is assessing the value of this trait in identifying
drought-adapted groundnut germplasm in large-scale breeding programme
(Wright, 1993).
Lawn (1989) argued that simple procedures should be developed to
identify sources for the putative traits, followed by divergent selection for
and against the trait in breeding programmes. Crop models will have to be
further developed to assess the value of individual traits in contributing to a
genotype's performance in a given drought environment (Muchow et ai.,
1991; Shorter et ai., 1991).
9.7.3 Drought management options
(a) Supplementary irrigation

Irrigation is a popular production practice wherever groundnut is culti-
vated under high input conditions. Pod yields as high as 10 t/ha have been
achieved under large-scale commercial conditions where irrigation has
been employed (Hildebrand, 1980), reflecting the high yield potential of
the crop. Various aspects of irrigation management, including irrigation
scheduling, have been described in detail by Boote et ai. (1982) and Boote
and Ketring (1990). In regions where the crop is grown under rainfed
conditions, supplementary irrigation is available only on a fraction of the
arable land. Where supplementary irrigation facilities are available, irriga-
tion water must be used efficiently to ensure the best returns to the farmer.
Increases in frequency or intensity of droughts, declining water tables and
increased costs of irrigation have led to considerable research effort on
irrigation management in the groundnut crop.
Variable results on the response of groundnuts to irrigation have been
obtained, depending upon several factors including the timing and amount
of irrigation, method of irrigation, the intensity of water deficit experi-
enced by the crop, and climatic conditions. Nageswara Rao et ai. (1985)
observed that drought during the pre-flowering phase, followed by ade-
quate water availability, resulted in pod yields of between 13% and 19%
greater than fully irrigated crops in a two-year study. This study showed a
significant interaction between ET and pod yield with different timing of
drought. The yield advantage in the pre-flowering drought treatment
occurred due to the better synchrony in pod set and pod filling following
release from the drought, which in turn resulted in an increase in the
number of mature pods at harvest (Nageswara Rao et ai., 1988). Several
reports have indicated that the pod-filling phase is most sensitive to
drought (Boote et ai., 1976; Pallas et ai., 1979; Nageswara Rao et ai.,
1985).
Effects of irrigation timing and intensity on yield of ground nut have been
examined in 22 groundnut genotypes by subjecting them to 12 different
types of single and multiple droughts (Table 9.4). As mentioned earlier,
the sensitivity of the crop to drought was less in early and mid-season
0
.,,,,
'"6
'"0
a5
-2
""" .
"",.
:s:
~
E
-4
."""
"
:g
""
~
"0
0
a.
-6
"" , .
"
"
-8
""
"
•
o 20 40 60 80 100
Effective duration of stress (days)
Figure 9.11 The effect of (.) drought or (.) non-limiting water during the
preftowering stage on the sensitivity of groundnut (mean of 22 genotypes) to
subsequent droughts of varying duration, ICRISAT Center, post-rainy season,
1982-83. (After Nageswara Rao and Williams, 1984.)
drought patterns compared with late-season droughts. However, it is

interesting to note that Nageswara Rao and Williams (1984) found that
exposing the crop to a short drought during the early vegetative phase
reduced the impact of a second drought at the seedling phase, indicating an
adaptive response of groundnuts to drought (Figure 9.11). It is possible
that early drought may enhance root development and reduce transpira-
tionallosses by limiting leaf area development, which subsequently allows
the use of soil moisture from deeper in the soil profile.
The beneficial effect of subjecting the crop to an early drought has
considerable practical implications for irrigation management. Since the
crop can endure long droughts during the early growth phase without
major yield losses, supplementary irrigation, when available, should be
used largely to support crop growth during the reproductiv-e stage. For
instance, in a five-year study in India, sowing of groundnut with one
supplementary irrigation prior to the arrival of the monsoon (in July)
resulted in moderate crop water deficits during the early growth stage, and
an increase in yield of up to 22% (Pasricha et al., 1987).
(b) Adaptation of genotypes to drought -prone environment

Manipulation of crop phenology to match the duration of soil water
availability is one of the most important ways of influencing crop perform-
ance in a given environment. One can match crop phenology to the
environment by either manipulating sowing date or selecting genotypes
with appropriate phenology to suit the environment.
Matching the environment to crop phenology

Selection of appropriate sowing dates to match crop growth with soil water
availability is an important management practice where groundnut
depends on seasonal rainfall (e.g. in India and Africa). Yield and phenolo-
gical traits vary significantly with changes in sowing dates, which can
influence soil and environmental components (Murthy and Nageswara
Rao, 1986; Kvien and Bergmark, 1987; Tsai etal., 1987; Bell, 1986; Bell et
al., 1991a). Several studies in India have shown that yields of groundnut
grown during the rainfed season (July-October) decline as the sowing date
is delayed beyond the end of July (Murthy and Nageswara Rao, 1986;
Hosmani et al., 1989; Dhoble et al., 1990). Here, the rainy season in most
areas ends by October, so that sowing beyond July exposes the crop to
terminal drought conditions.
Matching the genotype to the environment

Significant diversity exists in the groundnut germplasm for various pheno-
logical traits (Wynne and Coffelt, 1982) and offers the possibility of
selecting genotypes with desired phenology for use in breeding pro-
grammes. In regions where the growing season is longer. cultivars belong
to the virginia botanical group (subspecies hypogaea) are generally
selected; in regions where the season is shorter, spanish and valencia types
(subspecies Jastigiata) are selected. With perceivable changes in global
climate (principally temperature and rainfall patterns), it may become
necessary to match genotype more carefully to the length of growing
season. For example, groundnut production in Nigeria has declined
remarkably over the past few years due to severe droughts. This has
prompted the need for revision of recommendations on longer season
genotypes (Harkness and Dadirep, 1978, cited by Gibbons, 1978).
Agroclimatological analysis of major rainfed groundnut-growing environ-
ments in the semi-arid tropics (SAT) clearly indicates that growing areas in
the SAT are characterized by short growing seasons, i.e. 75-110 days
(Virmani and Piara Singh, 1986). This explains why short duration geno-
types are generally successful in West African regions (Gautreau, 1967;
Bocklee-Morvan, 1983) as well as in some parts of India (S.N. Nigam,
Principal Groundnut Breeder, ICRISAT, personal communication).
(c) Manipulation of plant density

Adjustment of plant population to match crop water demand to soil water
availability is another management option for growing groundnuts under
rainfed conditions. It is well known that dry matter accumulation under
400
E 300
I
~ •
"C
o
J§ 200
~ o
o
•
o 'A' spacing (35xl0 cm)
• '8' spacing (70xl0 cm)
100 o
o 'C' spacing (120xl0 em)
o
40 60 80 100 120
Transpiration (mm)
Figure 9.12 Relationship between transpiration and total dry matter of groundnut
grown at three plant densities under drought conditions, ICRISAT Center, post-
rainy season, 1981-82. (After Azam-Ali et al., 1989.)
non-limiting conditions is proportional to the amount of radiation intercep-

tion by the foliage (Biscoe and Gallagher, 1978; Azam-Ali et ai., 1989).
Similarly, Azam-Ali et al. (1989) have shown that the dry matter produced
is also tightly linked with crop transpiration (Figure 9.12). The tight
linkage between transpiration and dry matter production indicates that
yield cannot be enhanced by limiting transpiration, and hence that any trait
or management practice which improves the transpirational component of
total ET should improve yields. Obviously, manipulation of plant density is
an important way of optimizing leaf area to water supply and environ-
mental demand.
The relationship between groundnut productivity and plant density is
complex, particularly under water-limited conditions (Jones, 1986). The
optimum plant density under variable water availability will involve a
delicate compromise between maximizing water extraction and reducing
soil evaporation on the one hand, and conserving soil water for use during
pod filling on the other. Both the timing and severity of crop water deficits
occurring in response to rainfall availability and distribution, and soil water
holding capacity, will interact to determine the plant population/yield
response. For instance, groundnut crops grown at high plant popUlation
under severe end-of-season drought stress depleted the soil water rapidly
and led to severe crop water deficits during reproductive growth, and
reduced pod yield, compared with low density crops (Wright and Bell,
1992b). Indeed, the optimum plant density for groundnut has been shown
to decline as the severity of drought increases (Wright and Bell, 1992a):
while pod yield was maximized at over 100000 plants/ha under full irriga-
tion, only 40000 plants/ha were needed under a terminal drought stress.
These results indicate that significant savings in seed costs, and hence
increased profitability, could be achieved by reductions in plant population
under specific drought regimes.
Much of the information on plant density response in groundnut is
empirically based, and few attempts have been made to understand the
physiological factors responsible for the variation often observed in differ-
ent genotypes, drought patterns, soil types and localities. Nageswara Rao
et ai. (1989b) studied root and shoot growth in response to varying plant
density for groundnut (cv. TMV-2) grown on stored soil water. They found
that total weight and length of roots per unit area increased with plant
density, but proportional increases in shoot weight were greater, such
that the root to shoot ratio increased from 0.3 at high plant population
(35 x 10 cm spacing) to 0.5 under low plant popUlation (120 x 10 cm
spacing) (Table 9.6). The influence of plant population on the relative size
of roots and shoots has important implications for the water relations of the
crop in a given environment.
There have so far been limited efforts at predicting the plant population
requirements from knowledge of weather and soil parameters. Climatic,
soil and management factors, and the interaction between supply and
demand for water, determine the optimum plant population at which
maximum productivity can be obtained. Azam-Ali et ai. (1993) reported
that the optimum plant population requirements for various climates can
be determined using environmental features such as seasonal soil water
supply, saturation vapour pressure deficit, and crop parameters such as
water-use efficiency and harvest index. Bell et ai. (1991b) developed a
simple model, originally proposed by Gardner and Gardner (1983), for
predicting the relationship between groundnut pod yield and plant density
under non-limiting water conditions. The model provided a useful method
for summarizing key components of the pod yield/plant density relation-
ship, and served to highlight important characteristics associated with
different density responses among cultivars. Wright and Bell (1992a) ex-
tended this approach to characterize the relationship under water-limited
conditions. Some of the assumptions made in the model were shown to be
invalid under drought conditions; however, it was suggested that these
crop constants may be able to be related to either crop or soil water status,
in order to improve the predictive capability of the model under water-
limited conditions. The model provides a useful analytical tool for inter-
cepting groundnut plant density responses, by defining key physiologically
meaningful parameters. It has also been used recently to interpret the
differing ground nut population response observed between subtropical
and tropical regions (Wright, 1992).
TABLE 9.6 The mean weights of shoots (including pods) and the total weight and
length of the root systems per unit land area (from Nageswara Rao et aI., 1989b)
Spacing ++ Days Shoot plus Root weight Root length Root:

after pod weight Wa la total weight
sowing (glm2! (glm2) (kmlm2) ratio
A 47 49
A 60 134 58 3.8 0.30
A 76 206 68 4.7 0.24
A 90 267 143 6.6 0.36
A 97 230
B 47 30
B 60 68 41 2.7 0.38
B 76 113 58 3.7 0.33
B 90 155 120 5.6 0.43
B 97 147
C 47 16
C 60 44 44 2.8 0.49
C 76 71 49 3.2 0.41
C 90 105 101 4.6 0.49
C 97 100
Analysis of variance
Main effects
Spacing *** ** ** ***
Date *** *** *** **
Interactions
Spacing x date *** NS NS NS
*** and ** denote significance at the 1 and 2% levels, respectively:

'NS' dcnotes lack of significance at the 5% level.
Spacing: A = 35 x 10 cm; B = 70 x 10 cm; C = 120 x 10 cm.
(d) Intercropping
Intercropping of groundnut with various legume and cereal crops is a
popular practice, particularly under rainfed conditions where farmers have
limited access to irrigation. Many studies have reported advantages due to
intercropping in terms of land resource use, cost benefit ratio, and sustain-
able yields (Willey, 1981). Yield advantages in an intercropping system can
occur if the component crops have variable phenology and hence differing
resource requirements (primarily light and water) in space and time, The
extent of yield advantage, however, depends on the environmental re-
source capture, resource use and the resource conversion efficiency. There
are several reports on yield advantages of intercropping groundnut with
cereals (Reddy and Willey, 1981; Harris et ai" 1987), although the factors
causing yield advantage varied depending on the situation. In the inter-
cropping of sorghum with groundnut, shading of groundnut by sorghum
ameliorated the effects of high temperature and water deficit to a certain
extent, especially under drought (Harris and Natarajan, 1987). It was
suggested that alleviation of high temperature effects on flower production
resulted in a greater proportion of pegs forming pods in the intercropped
groundnut compared with the sole crop, leading to the observed increase in
harvest index of groundnuts. Despite a serious drought in 1984 in the
savannah zone of the Sudan, Arya et al. (1988) reported that intercropping
of millet with groundnut reduced soil loss due to wind erosion, and
increased yields compared with the sole-cropped groundnuts.
The significant variability in phenology and duration among different
botanical groups of ground nut genotypes makes it possible to intercrop
short and long duration genotypes (Nageswara Rao et al., 1990). Experi-
ments on intercropping of these in a ratio of 1:1 under a range of water-
deficit conditions imposed during the pod-filling phase resulted in an
increase in land equivalent ratio (LER) of up to 1.25. However, it was
interesting to note that specific combinations of cultivars were necessary to
maximize LER. Although the intercropping of early and late maturing
genotypes resulted in absolute benefits for total dry matter, there was no
apparent advantage in pod yields because of low partitioning in the case of
the long duration component genotype used in the study. The total yields
of this system could probably be improved by using a long season genotype
with greater partitioning of dry matter to pods. These results suggest that
there may be scope for achieving greater productivity in environments with
variable season length by intercropping late and early maturing groundnut
cultivars. Further research into the factors limiting exploitation of water by
the long season component of the intercrop following harvest of the early
genotype is needed to optimize the temporal and spatial utilization of light
and water resources. Such an option may be advantageous in regions
where hand harvesting is practised but may present management problems
in sowing and harvesting operations where mechanical cultivation is
practised.
(e) Mulching
As mentioned earlier, the linear relation between dry matter production
and amount of water transpired by the crop (Azam-Ali et al., 1989) means
that any practice or trait which increases transpiration can thereby increase
dry matter production. Under water-limited conditions, another manage-
ment option to enhance a crop's productivity is to increase the amount of
transpirable water, by using soil water conservation practices to reduce the
soil evaporation component of total evapotranspiration. One of the most
popular methods for reducing soil evaporation is by the use of soil mulches.
Araya et al. (1988) reported that, despite serious drought in the 1984
cropping season, mulching with surface residues and no till planting
increased soil moisture, biomass and seed yields on the cracking vertisols
and stabilized sands in the savanna zones of the Sudan.
Plastic mulches have been widely adopted in China for production of
groundnut under dryland conditions (Wang and Han, 1990). Besides
causing increased surface soil temperatures, which allows earlier sowing
times, the plastic mulch significantly reduces soil evaporative losses. For
instance, soil water content in the surface 10 cm was up to 5% higher
during early seedling growth under plastic mulch compared with uncovered
soil. Pod yield responses of greater than 50% have been recorded using
plastic mulch under rainfed conditions in Chinese groundnut production
systems.
9.8 CONCLUSIONS
Drought is recognized as one of the major constraints limiting groundnut

productivity in rainfed production areas. Plant water deficits strongly
influence physiological processes such as photosynthesis, stomatal conduc-
tance, leaf expansion and nitrogen fixation, which all ultimately have an
impact on pod yield performance. Groundnut is capable of withstanding
extremely low relative leaf water content (as low as 29%) and low leaf
water potential (as low as -10.0 MPa), indicating substantial dehydration
tolerance. The maintenance of turgor pressure through osmotic adjustment
has been demonstrated in groundnut leaves and pegs. However, the situ-
ation regarding the importance, extent and possible cultivar variation in
osmotic adjustment in groundnut is unclear.
Water deficits strongly influence root and shoot growth in groundnut.
Cultivar differences in root growth and soil water extraction capacity under
drought stress have been shown to exist. Further research effort is needed
to show that higher yield and improved drought tolerance are associated
with larger and deeper root systems. Simple, rapid and inexpensive
methods to screen for rooting characters which correlate with observed
field performance are needed. Pod yields can be severely reduced by water
deficits occurring during the pegging and pod set phase. Soil water content
can influence peg and pod development through direct effects on plant
water status and assimilate supply, or through effects on peg penetration
and calcium uptake in dry surface soil. Peg entry and subsequent pod
development have also been shown to be extremely sensitive to soil
hardness, which commonly increases with declining soil water content. The
partitioning of biomass to pods can vary widely, depending on the timing
and severity of water deficits, and strongly influences final pod yield. Both
genetic and environmental factors influence partitioning patterns in
groundnut.
There are a number of management and genetic options available to
References 325
ameliorate the effects of water deficits in groundnuts. Management op-
tions, such as supplementary irrigation and manipulating crop phenology
(by cultivar selection and/or sowing date) to match the duration of soil
water availability, are widely employed in many groundnut producing
regions. Current research has also shown that plant density manipulations
and intercropping of groundnut with other crops (or with different matur-
ity cultivars) can ameliorate the detrimental effects of drought by delaying
the onset of severe crop water deficits.
The genetic solution to drought is considered the best long-term strategy
to improve pod yield performance in water-limited environments.
Systematic attempts to screen groundnut genotypes for drought tolerance
have only been made at IRHO (Senegal) and ICRISAT (India). In these
breeding programmes, adapted genotypes have been selected solely on the
basis of increased pod yield and stability in drought-prone environments.
Future genotypic selection based on desirable drought tolerance traits
contributing directly or indirectly to superior performance under drought
conditions might be possible. For instance, current research is assessing the
potential of traits such as transpiration efficiency, partitioning of dry
matter to pods, and efficient root systems to improve the efficiency
of selection of drought-adapted germplasm in groundnut breeding
programmes.
ACKNOWLEDGEMENTS
We thank the Queensland Department of Primary Industries (QDPI) the

International Crops Research Institute for the Semi-Arid Tropics (ICRIS-
T AT) and the Australian Centre for International Agricultural Research
(ACIAR) for their support during the production of this chapter, which
was submitted as ICRISAT journal article No. JA 1457.
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CHAPTER 10
Diseases
K.J. Middleton, S. Pande, S.B. Sharma and
D.H. Smith
A well-balanced groundnut research team will always include a plant

pathologist, such is the importance of diseases in the crop. However, the
importance of a particular disease in a crop depends on a number of
environmental factors. Under cool wet conditions, sclerotinia blight and
web blotch may be important; bacterial wilt will be serious only under hot
wet conditions. The leaf spot diseases and nematodes are important when
the crop is grown regularly on the same soil; rust can be important under
drier conditions than those favouring leaf spots. Aflatoxin contamination is
worst under drought conditions, while those virus diseases spread by
aphids will be severe when groundnuts are growing well but other crops in
the area are suffering from water shortage. Other viruses which frequently
infect weed hosts are more serious when those weeds are prevalent in the
crop or its surroundings. The damping off of seedlings is most serious when
poor quality seed is planted without fungicidal protection. Under most
conditions, some incidence of disease can be expected, and in most pro-
duction areas disease control requires research input.
In this chapter, it is assumed that the reader knows the identity of the
disease causing concern and is consulting this volume to obtain more
information about it. Sometimes, the disease has been well covered in
another treatise, and in these cases the reader will be referred to the other
sources of information, but all diseases covered here are treated in ade-
quate detail to suit most purposes. To simplify discussion on the inter-
relationships between similar diseases, the chapter is organized so that
similar diseases are in proximity.
10.1 DISEASES CAUSED BY BACTERIA
10.1.1 Bacterial wilt

Bacterial wilt is caused by Pseudomonas solanacearum (E.F. Smith), an
aerobic Gram-negative rod shaped bacterium (Hayward, 1964). Virulent
Diseases caused by bacteria 337
isolates are fluidal, while avirulent isolates are butyrous in nature (Porter et
al., 1984; Cook et al., 1989). This is the only important bacterial pathogen
of groundnut. When this disease is severe, infection causes such rapid
wilting of stems and foliage that the affected leaves remain green. When
damage is less, due to lower disease pressure, the decline of the affected
plants is slower, resulting in general chlorosis. Entire plants or individual
branches may be affected and the plants often die, although recovery from
the wilted state is possible. The root system of infected plants may show
external and internal discoloration. The internal discoloration usually
extends into the lower stems. This symptom coupled with visible oozing
or streaming of bacteria from stems placed in water are diagnostic
characteristics.
Yield losses are closely related to plant death; quality is rarely affected
as the diseased plants generally have zero yield. In southern China, up to
10% of the planted area can experience yield losses of 30% , with extremes
of 80% loss (Sun et al., 1981). The groundnut crop grown on land not used
for paddy rice production is most seriously affected. In Indonesia, up to
25% of the yield of susceptible cultivars can be lost, particularly in South
Sumatra, West Java and South Sulawesi (Machmud, 1986). In Uganda,
yield losses of 24% have been recorded (Opio and Busolo-Bulafu, 1990).
This disease has also been recorded causing some crop loss in Mauritius
(Felix, 1971), Zimbabwe, Malaysia (Hayward, 1986), USA (Jenkins et al.,
1966) and recently in the Philippines (Natural et al., 1988), Fiji (Iqbal and
Kumar, 1986), Sri Lanka (Jayasena et al., 1988), Papua New Guinea
(Tomlinson and Mogistein, 1989) and India (Devi and Menon, 1979).
P. solanacearum has a wide host range of more than 30 families including
Solanaceae, Musaceae, Asteraceae and Fabaceae (Kelman, 1953). Both
weed and crop species are recorded as hosts, although some may not
always express symptoms. There are considerable differences in the host
range of the three recognized races of the pathogen: race 1 has a wide host
range, race 2 is restricted to Musa and a few related perennial hosts, and
race 3 is restricted to potato and a small number of other alternative hosts
(Buddenhagen and Kelman, 1964). A fourth race attacking mulberry has
been reported (He et al., 1983). In addition, four biovars have been
delineated by biochemical tests (Hayward, 1990). Groundnuts are nor-
mally attacked by biovar 3, occasionally by biovar 4, and by biovar 1 only
in the USA.
The pathogen can be introduced into clean fields through the use of
infected planting material of alternative crops, particularly vegetatively
propagated crops such as cassava. Contaminated soil and water could
introduce the organism, but this method of dissemination is not considered
to be important (Persley, 1986). Seed transmission was implied by Palm
(1922) but has not been reliably demonstrated, although Machmud and
Middleton (1990) produced sufficient evidence to urge caution and further
research in this important area.
338 Diseases
Pathogen survival is usually enhanced in wet, heavy clay soils, especially

where the crop is grown continuously (Sun et al., 1981) but may also be
high in sandy soil (He, 1990). Young plants are more rapidly diseased than
older ones (Miller and Harvey, 1932) and high soil temperatures favour
disease development (He, 1990). Low soil temperature favours survival in
soil (Akiew, 1986). High levels of soil organic matter may lead to a decline
in population of some races. Persistence between susceptible crops is
enhanced by the presence of weed hosts, which are common under the
environmental conditions and farming systems in those areas of the world
where bacterial wilt is an important groundnut disease (Yeh, 1990). It is
considered that some form of root injury may increase infection. When the
infected plants suffer water stress, the extent of infection rapidly becomes
apparent.
Crop rotations provide some measure of control, especially when the
rotations include a paddy rice phase (Kelman and Cook, 1977). As the
pathogen has a wide host range, the effect of rotations on disease incidence
is likely to be due to the management conditions imposed when different
crops are grown, rather than the presence of hosts more or less favourable
to pathogen persistence. Obviously, continuous cropping to groundnuts
will increase the disease problem.
It was with groundnuts that host resistance against P. solanacearum was
first successful (Porter et al., 1984) The resistance in Indonesian cultivar
Schwarz 21 has continued to be useful for almost 60 years, with no
apparent change in virulence of the pathogen. Cultivars developed in
China have greater resistance than Schwarz 21 (Yeh Wei-Lin, personal
communication) and Chinese breeding programmes aimed at bacterial wilt
resistance are better developed than any other (Boshou et al., 1990; Tan
and Boshou, 1990). Host plant resistance has also been used as a control
strategy in Uganda (Simbwa-Bunnya, 1972). However, resistance can be
eroded in a breeding programme unless efforts to test progeny against field
populations of the pathogen are made regularly. Most groundnut cultivars
developed in production areas not affected by bacterial wilt are susceptible
to the disease.
Biological control using rapidly growing bacteria has reduced the
incidence of bacterial wilt in other crops, e.g. potatoes, but control in
groundnuts using these methods has been attempted only recently
(He, 1990).
10.1.2 Bacterial leaf spot

It was stated in the previous section that bacterial wilt is the only important
bacterial disease of groundnut. While that statement is true, a symptom
commonly seen on groundnut leaves has recently been attributed to a
bacterial pathogen. Many workers have seen a characteristic leaf spot,
usually when the crop is growing well, often when the crop is young.
Diseases caused by viruses and mycoplasmas 339
The symptoms have been described as 'eye spot', due to the presence of a
small (up to 5 mm) tan spot on the upper surface of the leaflet, partly
surrounded by one or two semi-circular, lens-shaped tan lines. These may
eventually surround the central spot. Symptoms on the lower surface are
less distinct than for either of the two fungal leaf spots. In the past, this
symptom has been attributed to the effect of atmospheric pollutants, but
Joshi (pers. comm.) has confirmed that this disease is caused by
Xanthomonas sp.
The disease does not cause serious losses in the groundnut crop and it is
included in this chapter solely to assist in removing past confusion about its
cause.
10.2 DISEASES CAUSED BY VIRUSES AND MYCOPLASMAS
10.2.1 Peanut mottle virus (PMV)

First reported in USA (Kuhn, 1965), this potyvirus is commonly found in
groundnut production areas of eastern USA, South America, East Africa,
Asia and Australia. It is sometimes referred to under the synonyms
groundnut mottle virus, peanut mild mosaic virus or peanut severe mosaic
virus (Brunt et al., 1990). These names are evidence of the diversity of
symptoms produced by PMV in groundnuts. Most of these symptoms are
systemic mottling and mosaic of the foliage, with occasional necrosis. In
other hosts, however, the virus can cause more serious symptoms and can
be lethal (Behncken, 1970; Bock and Kuhn, 1975). Occasionally, the
symptoms can be so inconsequential (for example, wavy margins of leaf-
lets) that even experienced groundnut virologists have difficulty in detect-
ing them (Kuhn and Demski, 1984).
The natural host range of PMV includes a number of other legume crops
such as peas (Pisum sativum) , beans (Phaseolus vulgaris), lupins (Lupinus
spp.) and soybeans (Glycine max). These alternative crop hosts have
implications for control of the viru~ in groundnuts and for the effects of
groundnuts on other legumes in diverse cropping mixtures. Cassia spp. and
Vigna spp. are often infected, aiding in persistence of the virus within a
locality (Brunt et al., 1990).
The virus is naturally transmitted by aphids in a non-persistent manner,
although some vectors can remain infective for several hours (Paguio and
Kuhn, 1976). Seed transmission is also important (Kuhn, 1965) and is often
the means of introduction of the virus into a new production area, or a new
crop cycle in situations where groundnut production is seasonal. Seed
transmission rates vary from 0.02% to 2% in Vigna unguiculata (Demski et
al., 1983) and from 0% to 8.5% in groundnut (Reddy, 1991). PMV is also
seed transmitted in mung bean and cowpea but not in soybean and pea.
The effects of PMV on yield have been examined and reported on
several occasions. Abdelsalam et al. (1987) reported that the major effect
340 Diseases
of PMV infection was reduced seed size, not reduced seed number. Crop
losses are difficult to estimate, as yield responses depend upon a dynamic
interplay between the age of the plants when they become infected and the
ratio of infected plants to healthy ones. On an individual plant basis, yield
reduction of a plant infected within 5 weeks of sowing can be severe
(c. 50%) but the effect declines with delayed infection (Paguio and Kuhn,
1974). In general, it is estimated that crop losses due to PMV infection of
commercial crops are under 5% .
Control of seedborne viruses non-persistently transmitted by aphids is
difficult. Insecticidal control of the vector can be counterproductive, as the
presence of insecticide residues on the plants increases the feeding activity
of the vectors and hence the number of plants infected. Disease-free seed is
difficult to produce in commercial quantities when alternative sources of
the virus exist. Host-plant resistance remains the only economically and
environmentally sound approach to control.
Bijaisoradat et al. (1988) reported that none of 70 cultivars tested were
resistant to the four isolates of PMV used. However, low seed transmission
has been located among a few germplasm entries (Reddy, 1991) and could
be used to produce varieties which would no longer carry a reservoir of
virus into each field at planting. This avenue appears to be the only
possible option of control available. However, the apparent low yield
losses and the possibility that this form of resistance could be overcome by
aphid transmission from an external source has led to PMV resistance
being allotted a low priority in most breeding programmes.
For detailed information on the virus and its properties, see Brunt et al.
(1990).
10.2.2 Peanut stripe virus (PStV)

During the late 1980s, this potyvirus assumed the status of a major threat to
groundnut production in Asia and a potential threat to all production
areas. This is due to increased awareness of its distribution, the potentially
high crop losses which it may cause and its relatively high level of seed
transmission. Before the identity of the pathogen was clarified, it became
known under various synonyms including peanut mild mottle virus, peanut
chlorotic ring virus and sesame yellow mosaic virus. However, an inter-
national ad hoc committee published justification for adopting 'peanut
stripe' as the correct nomenclature (Demski et al., 1984a; Demski et al.,
1988).
The symptoms produced on groundnuts infected by PStV vary consider-
ably, depending on the strain of the virus (Demski et al., 1988). These
include mild mottle (produced by isolates from Thailand, China,
Indonesia, Myanmar, India, Philippines and USA), blotch (Thailand,
Philippines, USA and Japan), stripe (Thailand and Philippines), blotch-
stripe (Thailand and Indonesia), chlorotic ring mottle (Thailand), chlorotic
line pattern (USA) and necrotic (Thailand). This list does not suggest that
a particular strain occurs only in countries shown above for that symptom.
Limited investigations into the effect of symptom variants have been
undertaken (Rechcig et al., 1989).
The virus is widely distributed throughout Asia, from China to Indonesia
and Myanmar, and has been accidentally introduced to the USA and India
in recent years (Demski et al., 1984a, 1984b; Rao et al., 1988a, 1988b,
1989). It has not been reported from Africa or Australasia. The situation in
Papua New Guinea is unclear.
PStV is transmitted by seed, and consequently is of great concern in
those countries where it is currently not known. Reported transmission
rates vary from 0.3% (Saleh et al., 1989) and 8% (Xu et al., 1989) to 29%
(Rao et al., 1988b), depending on botanical type and cuItivar, and age of
the plants at the time of infection. The virus is efficiently transmitted by
several species of aphids in a non-persistent manner.
As well as groundnuts, lupins (Lupinus albus), soybeans, Dolichos
lablab, Vigna unguiculata and sesame are natural hosts of PStV and there
may be more as yet undetected (Wongkaew and Dollet, 1990).
Differentiation of PStV from PMV can be achieved through mechanical
inoculation of Chenopodium amaranticolor (PStV gives local lesions, PMV
does not infect), groundnut and Phaseolus vulgaris cv. Top Crop (PStV
does not infect, PMV gives local lesions). PStV infects P. vulgaris cv.
Bataaf and cv. Kintoki (Brunt et al., 1990; Demski et al., 1984a).
PStV is capable of causing serious losses. Although one set of experi-
ments (Lynch et al., 1988) showed no effect of the virus on growth, yield
and grade of Florunner, subsequent results showed that decreased nutri-
tive value and changed flavour characteristics may follow PStV infection
(Ross et al., 1989). Careful observation in South Sulawesi, Indonesia,
during the 1986 dry season (Middleton and Saleh, 1988) suggested yield
losses of 70% in individual plants, and almost all plants were infected. Xu
(1988) reported greenhouse tests showing 23% loss and estimated annual
production loss of 200000 tonnes in northern China. Adalia and Natural
(1988) reported up to 24% yield loss in the Philippines, depending on
cuItivar. Designed plant loss experiments (Saleh et al., 1989) have shown
that losses are dependent upon the age of plants when infected, with 50%
loss when plants are infected by 2 weeks after sowing (i.e. seed trans-
mission), 25% loss if plants are infected from 3 to 8 weeks of age, and 14%
loss if plants are infected between 9 and 10 weeks of age. In the same
report, crop losses of 10% to 40% were measured, again dependent upon
age at the time of infection. Wakman et al. (1989) reported yield losses of
15% to 28%. Wongkaew (1989) reported variation in measured crop losses
from year to year, also dependent on cultivars used. He showed, however,
that the 'chlorotic ring' isolate could reduce pod yield up to 42% in all
cuItivars. There can be no doubt that PStV can cause serious losses of
groundnut yield in Asian dry season production.
342 Diseases
However, control of non-persistently aphid-transmitted viruses which
have a wide host range is very difficult once they become endemic (Brunt,
1989). Quarantine restrictions are obviously important to limit the spread
of PStV into those countries where it has not been found. Imported
germ plasm can be non-destructively tested for the presence of PStV, using
serological techniques. Imported trade product from locations where PStV
is endemic should not be grown, either deliberately or by accident, as the
probabilities for vector transmission from seedlings are high.
Production of sufficient quantities of PStV-free seed is one possible way
of minimizing the effects of this virus, as seed transmission appears to be
the major source of infection - dispersal by aphids occurs over only limited
distances (Xu, 1988). However, there are great difficulties in producing
and distributing virus-free seed in countries where groundnut production is
normally conducted by many farmers each producing on a small scale.
Plastic film mulch, as used for groundnut production in China, appears
to reduce infections of isolated plots, possibly because of its insect repel-
lent action (Xu, personal communication). However, when used widely, it
may contribute towards a uniform high incidence of PStV in fields.
The most economic, reliable and environmentally acceptable method of
control is by the use of cultivars which are either immune or have high
levels of resistance to the virus and/or its vectors. The only immunity
located to date, despite testing of over 11 000 germplasm accessions from
ICRISA T's world collection, is in the wild species Arachis duranensis
(Saleh, personal communication) and in A. cardenasii (Rao et al., 1991)
after mechanical, aphid and graft inoculation. Several laboratories are
currently attempting to produce plants with high levels of resistance, using
genetic techniques.
Another avenue of genetic control is that of non-seed transmission, i.e.
to find cultivars in which the virus is not transmitted by seed. Genotypes
showing very little seed transmission have been identified (Zeyong et al.,
1989) and are being used in breeding programmes to develop cultivars with
good agronomic traits. The success of the non-seed transmission approach
depends upon the extent of virus infection in other hosts in the region and
the activity of insect vectors.
For detailed information on the virus and its properties, see Brunt et al.
(1990).
10.2.3 Peanut stunt virus (PSV)

First reported in USA (Miller and Troutman, 1966), this virus is widely
distributed in the south-eastern USA, east and south-east Asia, Europe
and Africa (Reddy, 1991). This cucumovirus causes severe stunting and
dwarfing of plant parts in the USA but does not cause severe stunting in
China (Zeyong et al., 1989). Leaves are malformed; infected plants are
chlorotic and small; malformed pods with a split pericarp wall are
produced. PSV occurs mainly in beans and forage legumes. Host range
includes a number of legume crops and plant species in the Cheno-
podiaceae, Compositae, Cucurbitaceae and Solanaceae (Fischer and
Lockhart, 1978).
The virus is transmitted in a non-persistent manner by aphids (Hebert,
1967). It is readily transmitted by sap inoculation and is also transmissible
by dodder (Miller and Troutman, 1966). Seed transmission in groundnut is
less than 0.1% (Troutman etal., 1967).
Control of PSV is difficult. There have been no reports of groundnut
genotypes with resistance to PSV (Tolin, 1984) but genotypes with less
severe symptoms are available (Culp and Troutman, 1968).
10.2.4 Peanut clump virus (PCV)

The appearance of stunted plants, often with mottling, mosaic or chlorotic
ring spot symptoms on foliage, in the same part of a field each time
groundnuts are grown suggests the presence of peanut clump virus. This
rod-shaped furovirus was first recorded from Burkina Faso, West Africa
(Thouvenel et al., 1974), and has since been recorded in Ivory Coast,
Niger, Senegal, South Africa and India (Brunt et al., 1990). The Indian
strain of PCV is sometimes referred to as Indian peanut clump virus
(IPCV).
The virus may be introduced into uninfested fields by sowing infected
seeds (Thouvenel and Fauquet, 1981). The seed transmission rate in
ground nuts may be 5-11 % and the virus is also seed transmitted (10%) in
Setaria italica, a symptomless host. After introduction, the virus is trans-
mitted to further groundnut plants by the soil-inhabiting fungus, Polymyxa
graminis. This accounts for the slow enlargement of affected areas and the
regular recurrence of the symptoms with each groundnut crop. Soil pH
affects transmission by P. graminis.
Other hosts of PCV include Sorghum arundinaceum, S. bicolor,
Pennisetum glaucum, Eleusine coracana (symptomless natural hosts);
Chenopodium amaranticolor (chlorotic local lesions, ringspots, and line
patterns extending along veins); wheat (systemic mosaic); Phaseolus vul-
garis, Vigna unguiculata, V. mungo (systemic ringspots and/or mosaic),
and Nicotiana benthamiana (systemic ringspots and leaf distortion).
The Indian isolates collected from various parts of that country produce
differing symptoms on experimental hosts, although they are indistinguish-
able on the basis of particle size and produce identical symptoms on
groundnut, mung bean (Vigna radiata) and Viciafaba. The hosts Canavalia
ensiformis and Nicotiana clevelandii x glutinosa hybrid were the most
useful for separating these isolates on the basis of symptom expression
(Nolt et al., 1988). Serological differences between IPCV isolates were
demonstrated.
The effects of PCV on yield depend on the age of the plant when
344 Diseases
infection occurs. Early infected plants produce a few small pods, and plants
infected later can suffer yield losses of 60% (Nolt and Reddy, 1984).
Control of this virus relies largely upon planting seeds from areas not
infested with PCV, although selection of planting date can allow crops to
escape infection (Nolt and Reddy, 1984). Some cultivar resistance has been
observed but screening for resistance should be done at a number of
different locations to assess reaction to more than one isolate (Nolt et al.,
1988).
Development of an improved PCV detection system is in progress
(D.V.R. Reddy, personal communication). For detailed information on
the virus and its properties, see Brunt et at. (1990).
10.2.5 Groundnut rosette virus (GRV)

GRV is one of the most important viruses on groundnut in Africa, where it
probably originated and to which it is apparently restricted. The virus
infection causes chlorotic rosette, mosaic rosette and green rosette symp-
toms (Reddy, 1991). Chlorotic rosette, characterized by severe stunting of
plants with isolated flecks or dark green coloured leaves, is more prevalent
in East and Central Africa. Mosaic rosette is recorded in East and Central
Africa: younger leaflets show conspicuous mosaic symptoms and stunting is
rather less pronounced than for chlorotic rosette. Green rosette, character-
ized by slight mottling of young leaflets and presence of yellow leaves with
green veins, occurs only in West Africa and Uganda. Plants infected with
chlorotic rosette bear smaller, curled and distorted leaflets; the stunting of
plants depends on time of the infection.
Rosette is caused by a complex of two viruses and a satellite RNA (Hull
and Adams, 1968; Dubern, 1980; Murant et at., 1988). GRV is mechan-
ically transmissible but requires the presence of groundnut rosette assistor
virus for aphid transmission (Hull and Adams, 1968). It is transmitted in a
persistent manner and over 92% transmission occurs within 10 minutes of
the beginning of the inoculation access period (Dubern, 1980).
High-yielding rosette resistant groundnut cultivars RMP 12, RMP 91
and KH-149A are available. Resistance is controlled by two independent
recessive genes and resistant genotypes are reported in Malawi, Nigeria
and West Africa (Berchoux, 1960; Gibbons, 1977; Harkness, 1977; Gillier,
1978). All available rosette resistant cultivars are late-maturing but early-
maturing ones are being developed. The disease is partially controlled by
destruction of volunteer ground nut plants, early planting, the use of high
plant density and the application of insecticides to control aphids.
For detailed information on GRV, see Reddy (1991).
10.2.6 Spotted wilt (tomato spotted wilt virus, TSWV)

The tomato spotted wilt virus has an extremely wide host range, including
many crops and weed species. It is occasionally known as pineapple yellow
spot virus. On groundnut, it can cause a wide range of symptoms (one

strain produces 'peanut yellow spot') and consequently is a virus that
requires careful diagnosis (Reddy et al., 1990).
For full coverage of possible symptoms, including colour plates, refer
to Reddy et al. (1990). The range of symptoms extends from general
chlorosis, through chlorotic mottles, lines or spots, to necrotic rings,
spots or streaks, and can lead to necrosis of the terminal apices
or buds - a symptom which has inspired adoption of the secondary
name 'bud necrosis' for the disease caused by TSWV. However, this
symptom does not occur in all situations and it is most common in India
and mid-western USA. In addition to the foliage symptoms, plants may be
stunted and axillary shoots may proliferate. Seed produced on plants
infected early in life can be shrivelled, and the testa is often discoloured red
or purple.
TSWV has spread worldwide (Brunt et al., 1990). Spotted wilt disease of
groundnuts was first reported from Brazil in 1941 (Costa, 1950) and
subsequently from South Africa (Dyer, 1949), Australia (Helms et al.,
1961), India, Nepal, Nigeria, People's Republic of China, Thailand and
USA (Reddy, 1984; Reddy et al., 1990).
This virus is not transmitted by seed, by plant-to-plant contact, or by
pollen. It can be mechanically inoculated into groundnuts but the main
method of transmission is by thrips. Transmission is persistent, i.e. an
insect, once it has fed upon an infected plant, retains the capacity to
transmit the virus throughout the remainder of its life. No helper virus is
required for successful transmission (Brunt et al., 1990).
Crops and weeds that are hosts of both the virus and its vectors and are
often found in association with groundnuts include Amaranthus spp.,
Datura stramonium, Glycine max, Helianthus annuus, Lactuca sativa,
Lycopersicon spp., Nicotiana tabacum, Phaseolus vulgaris, Solanum spp.,
Vicia faba and Vigna spp. (Brunt et al., 1990). Lycopersicon esculentum
produces systemic symptoms (Reddy, 1984). Diagnostic hosts are cowpea
(Vigna unguiculata) and petunia (Petunia hybrida). On cowpea cvs. C-152
or California Black Eye, TSWV produces concentric chlorotic and necrotic
lesions on leaves 4-5 days after inoculation, while necrotic lesions are
produced on petunia 3 or 4 days after inoculation (Reddy et al., 1990).
Precise confirmation of the presence of TSWV can be made with ELISA
procedures.
Serious yield losses from TSWV in groundnut have been reported from
India since the mid-1960s, from Australia since mid 1970s and from the
USA since the mid 1980s (Black, 1988). Of these three countries, the bud
necrosis phase is rare in Australia, and appears to be associated with high
temperature (Reddy et al., 1990).
Yield losses due to TSWV depend upon the incidence of infected plants,
their age when infected and environmental conditions. In India and the
USA, plants infected when under one month of age frequently die. TSWV
346 Diseases
is currently considered to be the most important virus in Indian groundnuts
(Amin, 1985).
Reducing losses due to TSWV in groundnuts depends on integration of
vector management, the virus and cropping practices. Control of vectors
with insecticides can be difficult as they habitually exist in parts of the plant
where insecticidal contact is difficult, and the species have high potential
for mass migration. However, use of systemic insecticides may be success-
ful in reducing TSWV effects, although unwanted side effects (such as
other insect problems) may follow. The vectors are parasitized at times and
use of insecticides could interfere with this process. Planting groundnuts at
a time to avoid migratory thrips can be most helpful, if practicable under
local conditions (Reddy et at., 1990).
Management of the virus inoculum is most readily achieved by elimin-
ation of weed hosts from the planting and its surroundings (Purss, 1962)
but intercropping with cereals can also be beneficial. Planting at a density
to give rapid canopy closure reduces the proportion of affected plants
(Reddy et at., 1990).
Host-plant resistance to TSWV has not been found, despite exhaustive
testing. However, 'field resistance' has been reported (Amin, 1985) and is
thought to be due to non-preference for particular cultivars by the vectors.
This aspect of control is being strenuously pursued.
For greater detail on the virus, its purification and antiserum production,
and on the vectors of TSWV, see Reddy et at. (1990).
10.2.7 Other viruses

Groundnut crinkle, groundnut eye spot, peanut green mosaic, passion fruit
woodiness, peanut veinal chlorosis, peanut yellow spot and big bud viruses
have been reported on groundnut. However, these do not seem to be of
major importance at present. For details, see Reddy (1991).
10.3 DISEASES OF FOLIAGE, CAUSED BY FUNGI
10.3.1 Early leaf spot

The presence of clearly defined brown lesions on the leaflets of groundnuts
suggests that one or both of the two common leaf spot diseases are
attacking the crop. These are among the most serious of groundnut dis-
eases when considered on a worldwide scale (Smith, 1984; McDonald et
at., 1985). It is common for both leaf spot diseases to occur simultaneously
on the same crop but this does not always occur.
Early leaf spot is caused by the fungus Cercospora arachidicota Hori.,
which has as its perfect state Mycosphaerella arachidis Deighton. The
symptoms of early leaf spot include circular or irregularly shaped lesions up
Diseases of foliage, caused by fungi 347
to 10 mm in diameter, although they may coalesce to form larger areas of
necrotic tissue, particularly when lesions include a major leaflet vein.
Necrosis extends to the lower surface of the leaflet. The lesions are
medium brown in colour on the upper surface and light brown on the lower
leaf surface. A chlorotic halo is often present around lesions but may be
absent under some environmental conditions or on some cultivars. Fungal
sporulation is sparse and difficult to see with the unaided eye; most occurs
randomly distributed on the upper surface (Smith, 1984; McDonald et al.,
1985). Early leaf spot is distinguished from late leaf spot by the lighter
colour of the lesion's lower surface and by the random, sparse distribution
of conidiophores on the upper lesion surface. Microscopic examination of
conidia clearly separates the two diseases, when C. arachidicola will be
found to produce subhyaline olivaceous conidia measuring 35-110 fLm X
3-6 fLm; they are much more slender than the conidia of the late leaf spot
pathogen, which are 20-70 fLm x 4-9 fLm and have a finely roughened wall
(Mulder and Holliday, 1974). The pathogen may also attack petioles,
stems and pegs (carpophores). In these instances, lesion shape is deter-
mined by that of the infected structure.
Early leaf spot is usually found wherever groundnuts are grown (Com-
monwealth Mycological Institute, 1966). It is common for both leaf spot
diseases to occur on the same crop with early leaf spot usually being
present before late leaf spot. Many factors determine whether early leaf
spot or late leaf spot predominates in a particular crop, but there appears
to be a tendency for late leaf spot to dominate over early leaf spot when
there has been a prolonged series of heavily diseased crops (McDonald et
al., 1985).
The damage caused depends upon cropping practices, rainfall and other
environmental variables and may vary from crop to crop. Loss of pro-
duction (or its equivalent in cost of control) from the combined effects of
the two leaf spot diseases can range from 10% to 50% (Smith, 1984;
McDonald et al., 1985).
C. arachidicola survives in crop debris from previously infected crops.
Seed transmission is unimportant, at worst being mere surface contami-
nation (McDonald et al., 1985). Conidia of C. arachidicola are dispersed
aerially from debris and infected crops, with rain increasing spore dispersal
(Smith and Crosby, 1973). No other hosts of C. arachidicola are known
(Smith, 1984). .
The perfect state of C. arachidicola is not uncommon but plays no vital
role in the development of disease epidemics. Infection is favoured by
temperatures of 25-30°C and high relative humidity (Smith, 1984;
McDonald et al., 1985). Early leaf spot develops under slightly lower
temperatures than does late leaf spot. Earliest infections are usually on
older leaves close to the soil surface (inoculum source) and, given con-
tinued environmental conditions suitable for infection, the epidemic will
continue until most of the foliage and stems are infected and defoliation
348 Diseases
occurs. As a result, pod yield and quality can suffer (Smith, 1984;
McDonald et ai., 1985).
The literature on control of the leaf spot diseases of groundnuts is vast,
and both diseases are controlled simultaneously by most control measures.
Biological control through the use of mycoparasites has been suggested but
not seriously pursued (McDonald et ai., 1985). Cultural practices which
may make a significant impact on the damage due to leaf spot include crop
rotation, destruction or removal of crop residues from previous groundnut
crops and controlled planting schedules - all designed to minimize the
exposure of the crop to inoculum at least until it has reached more
advanced stages of growth (Smith, 1984; McDonald et ai., 1985).
Fungicides have been developed for the control of leaf spot; they are
widely used in developed countries and with increasing frequency in devel-
oping countries. Most are applied as an aqueous foliar spray, although
dusting is still used in some situations. The earliest fungicides for this
purpose have a broad spectrum of biological activity but only provide
protection against infection. This means that it is necessary to apply them
before the crop is exposed to inoculum, under weather conditions suitable
for infection of the plant. Under most conditions suitable for groundnut
production, the period between fungal invasion and visible symptoms of
leaf spot is 8-10 days, and some modern fungicides have the capacity to
control infections after invasion, until the first symptoms are appearing.
Some of the earliest systemic fungicides effective for leaf spot control
affected the growth of pathogen in a very simple way and, after widespread
use, 'fungicide resistance' developed: the pathogen changed its physiology
sufficiently to be able to grow in the presence of the fungicide (Smith,
1984).
Some wild Arachis spp. are either immune or highly resistant to early
leaf spot (McDonald et at., 1985) and a long-term goal of many groundnut
breeding programmes is to include resistance to early leaf spot as a trait in
released lines. Progress is being made towards multiple resistances of a
level high enough to eliminate or significantly reduce the need for fungi-
cides in lines which are also of acceptable marketable type and which
produce high yields, but to date the use of fungicides has not been
superseded.
10.3.2 Late leaf spot

Late leaf spot, caused by the imperfect fungus Phaeoisariopsis personata
(Berk. & Curt.) V. Arx. [Cercosporidium personatum (Berk. & Curt.)
Deighton], is widely distributed throughout the world (Commonwealth
Mycological Institute, 1967). Late leaf spot along with early leaf spot
(Cercospora arachidicoia Hori.) can cause pod yield losses of up to 50%,
worldwide (McDonald and Fowler, 1977; Smith, 1984; McDonald et ai.,
1985). In India, leaf spots and rust (Puccinia arachidis Speg.) normally
occur together and yield losses as high as 70% have been attributed to their
combined attack (Subrahmanyam et al., 1980).
Symptoms of the disease have been described by several workers
(Woodroof, 1933; Jenkins, 1938; Jackson and Bell, 1969; Feakin, 1973;
Garren and Jackson, 1973; Porter et al., 1982, McDonald et al., 1985).
Recently Smith et al. (1992) reviewed this disease in greater detail. The
symptoms of the disease are influenced by host genotype and environ-
mental conditions. Late leaf spots are first recognizable as small necrotic
flecks that enlarge and become light to dark brown measuring from 1 to
10 mm in diameter. Lesions formed by P. personata tend to be smaller,
more nearly circular, and darker than those formed by the early leaf spot
pathogen. On the abaxial surface the lesions are black and slightly rough in
appearance. Late leaf spot fungus usually sporulates on the abaxial surface
of the leaflet and conidial tufts of P. personata are macroscopically visible
as raised circles. The yellow halo which is often present around early leaf
spots is often less conspicuous in or absent from late leaf spot lesions.
Although symptoms are useful diagnostic features, positive identification
of the late leaf spot requires microscopic examination of the conidiophores
and conidia. The late leaf spot pathogen also produces lesions on petioles,
stems, stipules and pegs in the later stage of an epidemic. These are oval to
elongate and have more distinct margins than the leaflet lesions.
The perfect state of the late leaf spot pathogen is Mycosphaerella
berkeleyii W.A. Jenkins. However, the nomenclature of the anamorph of
this fungus has gone through several changes in the literature. Until
recently the combination Cercosporidium personatum (Berk. & Curt.)
Deighton was commonly used. In 1983, Von Arx proposed the new
combination Phaeoisariopsis personata (Berk. & Curt.) v. Arx. and de-
scribed the pertinent morphological characters of the anamorph as follows:
stroma dense, pseudoparenchymatous, up to 13 f.Lm in diameter, conidio-
phores numerous, pale to olivaceous brown, smooth, 1-3 geniculate,
10-100 x 3.0-6.5 f.Lm in size, conidial scars conspicuous, prominently
2-3 f.Lm wide; conidia medium olivaceous, cylindrical, obclavate, usually
straight or slightly curved, wall usually finely roughened, rounded at the
apex, base shortly tapered with a conspicuous hilum, 1-9 septa not
constricted, mostly 3-4 septate, 20-70 x 4-9 f.Lm in size. Jenkins (1938)
described the teleomorph of the fungus.
Although the teleomorph of the late leaf spot pathogen is known, the
ascigerous and spermagonial states are not generally regarded as important
sources of primary inoculum. The principal source of initial inoculum is
probably conidia that are produced on groundnut crop residues in the soil.
Following early rains, conidia are blown or rain-splashed on the leaves
of young groundnut plants, where they initiate the disease cycle.
Temperatures in the range of 25-30°C and high relative humidity favour
infection and disease development (Lyle, 1964; Jensen and Boyle, 1965).
Lesions develop within 10-14 days of initial infection. Duration of leaf
350 Diseases
wetness is another important factor in late leaf spot infection. In favour-
able conditions, disease progress continues throughout the season and may
lead to total defoliation of the infected plants. Spatial and temporal spread
of late leaf spot have been studied by several workers (Venkataraman and
Kazi, 1979; Knudsen et al., 1987; Alderman et al., 1989) and used in
developing a computer simulation model to produce schedules for fungi-
cide spraying (Phipps and Powell, 1984; Smith, 1986; Knudsen et al., 1988;
Alderman et al., 1989). The pathogen may also survive from season to
season on volunteer groundnut plants and on 'ground-keepers' (Hem-
ingway, 1954). No authentic host species outside the genus Arachis are
reported. Long-distance dispersal of P. personata is not known. Heavy
application of nitrogen and phosphorus fertilizers increases severity of late
leaf spot (Shanta, 1960).
Late leaf spot can be partially controlled with crop management prac-
tices that reduce initial inoculum (Smith and Littrell, 1980; Smith, 1984).
Additionally, fungicidal control of the disease has proved effective and
economical. Large numbers of fungicides have been developed and exten-
sively evaluated both in developed and developing countries (McDonald
et al., 1985; Subrahmanyam and Ravindranath, 1988; Jackson and
Damicone, 1991). Possible effects of fungicides on non-target organisms
should be considered. Exclusive use of chlorothalonil on a calendar sched-
ule has been shown to increase sclerotinia blight (Porter, 1980a). Similarly,
higher levels of southern stem rot caused by Sclerotium rolfsii Sacco have
occurred in fields of Florunner groundnuts sprayed with benomyl (Back-
man etaZ., 1975, 1977).
Breeding resistant cultivars is one of the best means of reducing crop
yield losses from late leaf spot. Resistance levels are not great enough to
discontinue the use of foliar fungicides but several fungicide applications
may be eliminated when moderate levels of resistance are integrated with
chemical control (Shokes and Gorbet, 1991). The decision as to whether or
not disease control should be recommended has to be made at the local
level, as it takes several factors into consideration such as extent of loss,
the cost of control measures, and economic and other returns expected.
Several mycoparasites have been reported to parasitize the late leaf patho-
gen but no serious attempts have been made to use them in the integrated
management of leaf spots (Subrahmanyam and Ravindranath, 1988).
10.3.3 Rust
Rust caused by Puccinia arachidis Speg. is one of the most destructive
fungal foliar diseases of groundnuts in many groundnut-producing
countries of the world (Bromfield, 1971; Hammons, 1977; Subrahmanyam
and McDonald, 1983; Smith, 1984; McDonald and Subrahmanyam, 1992).
Yield losses from rust are over 50% (Subrahmanyam et aZ., 1980a; Ghuge
et aZ., 1981). The effect of rust on groundnut yields is likely to increase with
improvement in the production situation with more favourable environ-
mental conditions. However, damage increases with multiple pathosystems
particularly where rust and late leaf spot are involved (Savary and Zadoks,
1992a, 1992b).
Rust disease can be readily recognized by the appearance of orange-
coloured pustules or uredinia (uredia) on the abaxial surface of the ground-
nut leaves. Uredinia then rupture to expose masses of reddish-brown
urediniospores (uredospores), which land on the leaf surface. If tempera-
tures are in the 15-30°C range and leaves are wet with dew or rain, the
urediniospores germinate and produce appressoria and infection hyphae,
which penetrate the leaf through stomata (Subrahmanyam and McDonald,
1983; Butler and Jadhav, 1991). The uredinia'appear within 8-20 days but
chlorotic flecks are usually visible macroscopically 2 or 3 days before the
appearance of erumpent sori (pustules). Incubation period is greatly
influenced by environmental factors and host genotype (Mallaiah, 1976;
Sokhi and Jhooty, 1982; Subrahmanyam et ai., 1983). Pustules appear first
on the abaxial surface but in extended periods of high humidity are
sometimes formed on the adaxial surface. In highly susceptible cultivars,
the primary pustule is encircled by colonies of secondary pustules. The
pustules, which develop on all aerial plant parts except flowers and pegs,
are usually circular and range from 0.3 to 1.4 mm in diameter. In contrast
to rapid defoliation associated with leaf spots, leaves infected with rust
become necrotic and dried, and remain attached to the plants for several
days (Subrahmanyam and McDonald, 1983).
In general, rust fungi are known for their complex life-cycles. Some
species at times produce five different kinds of spores and require two
unrelated hosts to complete the life-cycle. However, groundnut rust is
known almost exclusively by its uredinial stage. Uredinia are predomi-
nantly located on the underside of the leaf, scattered or irregularly
grouped; they are round, ellipsoid or oblong, and dark cinnamon brown
when mature. The ruptured epidermis is conspicuous: urediniospores are
broadly ellipsoid or obovoid, 16-22 x 23-29,....m in size, brown-walled,
1-2.2 ,....m thick, and finely echinulate. They have usually two but occasion-
ally three or four germ pores, nearly equatorial, often in flattened areas.
A few reports of the occurrence of teliospores, mainly from South
America (Arthur, 1934; Jackson and BelI, 1969; Hennen et ai., 1976;
Cummins, 1978), suggest that telia chiefly occur on the underside of leaves;
they are 0.2-0.3 mm in diameter, scattered, prominent, naked and chest-
nut or cinnamon brown, becoming grey or almost black. Ruptured epider-
mis is prominent; teliospores are oblong, obovate, ellipsoidal or ovate with
a rounded to acute and thickened apex. They are constricted in the middle,
gradually attenuate at both ends, smooth walled, light yellow or golden
yellow to chestnut brown, predominantly two-celled, sometimes three or
four cells, 38--42 x 14-16,....m with walls 0.7-0.8,....m thick at the sides and
2.5--4.0,....m thick at the top. The apical thick part is almost hyaline. The
352 Diseases
pedicel is thin-walled, usually collapsed laterally, hyaline, up to 35-65 j.Lm
long and normally detached at the spore base. It germinates at maturity
without dormancy. Puccinia arachidis has not been observed to produce
metabasidia and basidiospores.
Rust disease is known to perpetuate, spread and cause severe epidemics
by means of urediniospores. There are a few records of the occurrence of
the telial stage on cultivated groundnuts in South America (Spegazzini
1884; Hennen et al., 1976) and on wild Arachis species (Guarch, 1941;
Bromfield, 1971). It is not known if the fungus produces spermatia and
aecia or if any alternate host is involved in the life-cycle. Subrahmanyam
and McDonald (1982) have examined various common crop and weed
plants as possible hosts for the groundnut rust pathogen, but no case of
infection has been recorded.
Urediniospores are short-lived in infected crop debris (Mallaiah and
Rao, 1979; Subrahmanyam and McDonald, 1982). It is therefore unlikely
that the fungus is perpetuated from season to season in crop debris under
hot climatic conditions when there is a break of over 4 weeks between crop
seasons (Subrahmanyam et al., 1980b). The pathogen may survive from
season to season on volunteer groundnut plants. Long-distance dissemina-
tion of the disease may occur via airborne urediniospores and/or by the
movement of infected crop debris (van Arsdel and Harrison, 1972). There
is no authenticated report of rust being internally seedborne or spread by
germplasm exchange.
Cultural practices, host-plant resistance and fungicide usage are inte-
grated into effective management strategies to minimize initial levels of
disease and slow down the progress of rust epidemics. Wherever possible,
successive groundnut crops should not be planted where viable uredinio-
spores are present. Volunteer groundnut plants and 'ground-keepers'
should be eradicated. If cropping systems permit, times of sowing should
be adjusted to avoid infection of the crop from outside and to avoid
environmental conditions conducive to rust build up. Strict plant quaran-
tine restrictions should be enforced to avoid the spread of rust conidia on
pods or seeds to areas where the disease is not present.
A number of fungicides developed for the control of leaf spots have been
evaluated for effectiveness against rust (Subrahmanyam and McDonald,
1983; Subrahmanyam et al., 1984). However, the old established Bordeaux
mixture has given useful control of both rust and leaf spots. Among the
new generation systemic fungicides, chlorothalonil gives good control of
both the diseases but calixin is effective only against rust, while benomyl is
effective against leaf spots but not against rust.
The rust resistance available in the cultivated groundnut is of the 'slow
rusting' type, i.e. resistant genotypes have increased incubation periods,
decreased infection frequency, reduced pustule size and spore production
and reduced spore viability (Subrahmanyam et al., 1980b; Subrahmanyam
and McDonald, 1983). When rust resistance is combined with resistance to
leaf spots and judicious use of fungicides, these diseases can be controlled
effectively and economically.
10.3.4 Web blotch

Web blotch was first recognized on dead groundnut leaves in Russia
(Woronichin, 1924). Similar disease symptoms were observed on ground-
nut leaves in Brazil (Cruz et al., 1962), Rhodesia (Rothwell, 1962),
Argentina (Frezzi, 1969), and the United States (Pettit et al., 1973). Since
then it has been reported from Canada, China and Australia (Taber,
1984). The disease is also known as Phoma leaf spot, Ascochyta leaf spot,
spatselviek, muddy spot and net blotch. It appears to be of increasing
importance in the United States (Phipps, 1985), and in South Africa it is
considered as one of the most important foliar diseases of groundnut. It has
been observed in groundnut-producing countries of Africa since 1946
(Taber, 1984).
Though the disease has been largely recognized to be caused by the
imperfect fungus Phoma arachidicola Marasas, Pauer, and Boerema,
generic distinctions between Phoma and Ascochyta and among Mycos-
phaerella, Didymosphaeria and Didymella need to be clarified before
assigning proper anamorphic and teleomorphic names (Taber et al., 1981).
Web blotch is characterized by a webbing or net-like pattern of tan to
bronze colour on the adaxial surface of the leaves. Another pertinent
symptom is the production of larger and nearly circular tan to dark brown
blotches on the adaxial leaf surface. Lesions coalesce to encompass the
whole leaf area. Rarely, a leaf flecking is observed. Older lesions become
dry and darker in colour than the younger ones. Lesions on the abaxial
surfaces are less prominent and pale brown in colour. The disease is
capable of causing severe defoliation, especially when associated with late
leaf spot.
Morphological features of P. arachidicola have been described in detail
by several researchers (Marasas et al., 1974; Taber et al., 1981). Pycnidia
which form on the infected host tissue, are 80-120/-Lm in diameter.
Pycnidial cell walls are translucent. The mycelium is 2-8.0 /-Lm in diameter,
olivaceous to brown and septate. It also forms dark chlamydospores (7-19
x 8 x 20 /-Lm), both aerial and submerged. Pycnidiospores are formed in
basipetal succession on short conidiogenous cells and are single-celled.
Pseudothecia have also been observed. They are dark brown, globose,
short-beaked to unbeaked, single, and 60-147 /-Lm in diameter. Asci (36-60
x 10-17 /-Lm) are bitunicate and cylindric to cylindric-clavate. Ascospores
(12.5-16 x 5.7/-Lm) are divided into two sections. Ascospores are two-
ceIled, smooth hyaline to pale yeIlow and become darker with age.
In 1975, Philley studied web blotch in detail and found that pycnidio-
spores, ascospores and chlamydospores are capable of initiating disease.
These inoculum propagules infect leaflets, petioles, stipules and stems.
354 Diseases
Germ tubes penetrate directly through the cuticle. Appressoria with small
infection pegs are formed near the germinated conidia. Epidermal pene-
tration is intercellular. The disease appears to be most severe during cool
seasons with high relative humidity. Groundnut is the only natural host.
Sweet clover and hairy vetch were found to be the most susceptible when
six leguminous genera were artificially inoculated (Philley, 1975). There is
evidence that airborne inoculum carried from South America during
Hurricane David introduced the disease to North Carolina and south-
eastern Virginia (Phipps, 1985).
Web blotch can be effectively controlled by commonly recommended
fungicides such as chlorothalonil and mancozeb plus benomyl. Crop ro-
tations and planting of less susceptible cultivars can decrease the incidence
of the disease (Phipps, 1985).
10.3.5 Pepper spot and leaf scorch

The pepper spot and leaf scorch disease of groundnuts caused by the
perfect fungus Leptosphaerulina crassiasca (Sechet) Jackson & Bell, has
been reported from Madagascar (Sechet, 1955), Taiwan (Yen et al., 1956),
India (Nayudu, 1963), Argentina (Frezzi, 1965) and USA (Luttrel and
Boyle, 1960; Pettit et al., 1968). Since then it has been reported from
Mauritius (Anon., 1960), Malawi (Mercer, 1977), Nigeria (McDonald,
1978) and Nepal (Pande et at., 1993). In comparison with early leaf spot,
late leaf spot and rust, this disease is considered of minor economic
importance throughout the groundnut-growing countries of the world
(Porter et al., 1982).
The disease is principally recognized by two types of symptoms. Pepper
spots are dark brown or black and restricted to the adaxial leaflet surface.
These lesions later enlarge and measure up to 1.0 mm in diameter. Lesions
are irregular to circular in shape and sometimes depressed. The leaf scorch
phase of the disease is commonly identified as wedge-shaped lesions with a
yellow zone adjacent to tips or margins of leaflets.
The causal agent of this disease is observed only in the sexual state
(Jackson and Bell, 1968). Pseudothecia (perithecia) are initially sub-
merged, later erumpent, amphigenous, yellowish-brown, spherical, thin-
walled, and 60-120 /Lm in diameter. The ostiole is short and papillate.
Pseudothecia contain 8-20 asci. Asci (50-80 x 25-55/Lm) are hyaline,
ovoid to broadly clavate, and have short stripes. Paraphyses are absent.
Each ascus contains eight ascospores, hyaline initially but becoming
yellowish-brown with age. Ascospores are oblong to ellipsoidal and
measure 23-40 x 11-17 /Lm, with transverse septa, and commonly con-
stricted at the septa.
The pathogen survives in necrotic leaflets and produces abundant asco-
spores. Ascospores are ejected up to 7.5 /Lm in experimental conditions
(Mallaiah and Rao, 1976). Peak dispersal periods of ascospores occur at
the end of the dew period and at the onset of rainfall (Smith and Crosby,
1973). Yen et al. (1956) observed that cardinal temperatures for in vitro
growth are 8, 28 and 35°C.
The fungus is known to infect only members of the genus Arachis. The
disease is apparently controlled by the fungicides commonly used to con-
trol early and late leaf spots (McGill and Samples, 1965). Host-plant
resistance is not available and most of the groundnut lines evaluated were
found to be susceptible (Porter et al., 1971).
10.3.6 Anthracnose
Several species of Colletotrichum have been reported to cause anthracnose
disease on groundnuts. However, C. mangenoti Ohevaugeon, C. arachidis
Sawada and C. dematium Pers. ex. Fr. Grove are the commonly associated
pathogens of Arachis spp. The disease has been reported on groundnuts
in the United States, Uganda, Taiwan, Senegal, India, Tanzania
(Tanganyika) and Argentina (Porter et al., 1982; Smith, 1984). At present
the disease is considered of minor importance throughout groundnut-
growing countries.
On plants infected with Colletrotichum mangenoti the leaflets, petioles
and stems display grey to brownish-grey lesions which are generally mar-
ginal to elongate (Chevaugeon, 1952). Plants infected with C. dematium
have small water-soaked yellow spots, 1-31 mm in diameter (Saksena et
ai., 1967). Symptoms induced by C. arachidis are scattered lesions
(3-5 mm), circular to irregular, with greyish-white centres surrounded by
brown borders (Sawada, 1959).
Morphological details and relationships of these three species of
Colletotrichum have been discussed in greater detail by several workers
(Frezzi, 1965; Jackson and Bell, 1969; Porter et ai., 1982) who have also
described the Colletotrichum species that infect wild groundnuts.
Very little is documented on disease management. Copper fungicides,
Bordeaux mixture (Chohan, 1974) and mancozeb (Singh et ai., 1975) have
been effective in controlling the disease.
10.3.7 Scab
Scab caused by Sphaceioma arachidis Bit. & Jenk. was first observed in
diseased groundnut plant materials collected in 1937 in Sao Paulo, Brazil
(Bitancourt and Jenkins, 1940). It has also been reported from Argentina
(Ojeda, 1966).
Primary symptoms appear as small spots (~1 mm diameter), round to
irregular in shape, on the adaxial leaflet surfaces. Spots are tan with
narrow, brown marginal lines. As the disease progresses, spots appear on
both sides of the leaf and become light tan with raised margins and sunken
centres on the upper leaf surface (Bitancourt and Jenkins, 1940; Giorda,
1984). On the petioles and stems, spots are larger and more irregular in
356 Diseases
appearance than on the leaves and become cankerous with age. In late
stages of disease development, lesions coalesce and the infected plants
grow in a sinuous fashion and become stunted. Some lesions acquire a
cork-like appearance and cover nearly all the plant parts, induding pegs.
The disease is endemic under both dry and humid conditions. However,
the fungus produces fruiting structures under conditions of high humidity
and these cover the lesions with an olivaceous velvety growth.
Host-plant resistance has been reported from Argentina (Ojeda, 1966)
and Brazil (Soave et al., 1973). The fungus persists in crop debris. Cultural
practices such as rotation may be useful in reducing the initial inoculum
and disease. Benomyl is found effective in controlling scab. Some evidence
suggests the seedborne and seed-transmitted nature of the disease (Giorda,
1984).
10.3.8 Pestalotiopsis leaf spot

A leaf spot of groundnut caused by Pestalotiopsis arachidis Satya was first
observed in India (Satya, 1964). Similarly, P. adjusta and P. vericolorhave
been reported to cause leaf spot diseases in Nigeria (McDonald, 1978).
Lesions caused by P. arachidis are dark brown and circular, and are
surrounded by a yellow halo. They are commonly restricted to either side
of the midrib. The centres of diseased leaf tissues are marked by the
presence of black, spherical acervuli. The fungus produces pink mycelium.
Conidia are ellipsoid-fusoid, four-celled and 17 x 6.9 fLm in size. At
present, this is not considered an important disease of the groundnut.
10.3.9 Phoma blight

Phoma blight, caused by Phoma microspora Balasubramanian and
N arayanasamy, was first observed in Tamil N adu in India (Balasubrama-
nian and Narayanasamy, 1980). Symptoms of the disease appear as scat-
tered light-brown lesions on the lower leaves of a crop more than 50 days
old. However, in artificial inoculations the fungus is capable of producing
typical lesions even on apical leaves, petioles and stem and can cause die-
back. Lesions on the stem are dark brown or greyish with irregular
margins. Numerous pycnidia have been observed on dead leaf tissues.
Currently the disease is restricted to Tamil Nadu, India, where it causes
considerable damage to the groundnut crop. Dhanju and Chohan (1974)
found P. glomerata associated with bud blight affected groundnut plants.
Little is known about the biology of these pathogens or the epidemiology
of the diseases caused by them.
10.3.10 Powdery mildew

Powdery mildew of groundnut, caused by Oidium arachidis Chorin, has
been reported from Mauritius, Portugal, Tanganyika and Israel (Porter
et al., 1982). Chorin (1961) named the powdery mildew fungus. Symptoms
of the disease are mainly restricted to the upper leaf surface. Oidia vary in
size and measure 31-44 X 13-15 /-Lm. Conidiophores bear one or two oidia
in dry weather, but chains of three or four oidia are common in un-
disturbed humid conditions. Subspherical pyriform haustoria develop in
epidermal cells (Chorin and Frank, 1966). In Israel, development of the
disease on groundnut foliage is rapid at 25°C.
10.3.11 Phyllosticta leaf spot

Leaf spots caused by Phyllosticta spp., though geographically widely dis-
tributed, are among the minor diseases of groundnut (Jackson and Bell,
1969). Disease has also been reported from India (Rao, 1963), Barbados
(Norse, 1974) and Nigeria (McDonald, 1978).
Frezzi (1960) described the symptoms of the disease as circular to oval
lesions of 1.5-5.0 mm in diameter with definite borders, halos absent,
reddish-brown on the perimeters and becoming lighter or tawny in the
centre. Shot-hole symptoms develop in some spots.
In India, Rao (1963) described Phyllosticta arachidis-hypogaea Vas ant
Rao as the cause of a leaf spot. P. sajoecola Massal was reported to cause a
similar leaf spot in Africa (Chevaugeon, 1952). The morphological descrip-
tion of these fungi is given in greater detail in the monograph by Jackson
and Bell (1969).
The fungus can remain viable in mycelial form for ~1 year on infected
crop debris and is the primary source of initial infection (Patil, 1981).
Optimum temperature and pH for spore germination and growth of
P. arachidis-hypogaea are 25-30 °C and 5.5-6.5, respectively (Patil, 1981).
Authenticated disease management practices are not known. However,
a few fungicides can inhibit the growth of the fungus in vitro (Patil,
1981).
10.3.12 Alternaria leaf spot

Leaf spot diseases of groundnut caused by both known and unknown
species of Alternaria have been reviewed recently by Subrahmanyam and
Ravindranath (1988), but currently most of them are regarded as minor
diseases worldwide. However, leaf spots caused by Alternaria arachidis
Kulk. and A. alternata (Fr.) Keissler are of growing importance on
the post-rainy season irrigated groundnut crop in southern India
(Subrahmanyam et al., 1981). Generally orange-brown necrotic spots in
the interveinal areas of leaves and extending into veins and vein lets are
characteristic of infection caused by A. alternata. The lower surface of the
lesions is sometimes covered by a light to heavy olive brown-green sporula-
tion. Porter et al. (1982) indicated that Alternaria spp. are frequent col-
onizers of necrotic groundnut leaf tissues. Several fungicides are being
358 Diseases
evaluated in various research centres in India (Subrahmanyam and
Ravindranath, 1988) and it appears that some modern fungicides have the
capacity to control alternaria leaf spot.
10.3.13 Zonate leaf spot

A minor foliar disease caused by Cristulariella moricola (Hino) Redhead
(Syn. C. pyramidalis Waterman and Marshall) was reported by Smith
(1972). The perfect state of the fungus is Sclerotium cinnamomi Sawada.
The disease is observed late in the growing season. Symptoms appear as
necrotic spots on the abaxial leaf surface. Lesions range from 2 to 12 mm in
diameter. Small lesions have a light brown centre surrounded by a brown-
ish ring of necrotic tissue and characteristic zonations are present on both
leaf surfaces. Pyramidal heads (conidia) form on both sides of the leaf.
They develop only on necrotic tissues, sometimes singly but rarely in
clusters of up to 45 per lesion. Sclerotia are produced on potato dextrose
agar and V8-juice agar.
10.3.14 Groundnut leaf blight

Subrahmanyam (1979) reported Myrothecium roridum Tode ex Fr. as the
causal agent of a leaf blight disease of groundnut in India. Leaf lesions are
round to irregular, 5-10 mm in diameter, grey in colour and surrounded by
a chlorotic halo. Diseased leaves have a blighted appearance after lesions
coalesce. Black fruiting bodies, often arranged in concentric rings, form on
the adaxial and abaxial leaf surface. Currently the disease is not economi-
cally important in the groundnut-producing countries of the world.
10.3.15 Melanosis
Melanosis of groundnut leaves caused by Stemphylium botryosum Wallr.
has been reported in Argentina (Frezzi, 1960). The symptoms of the
disease are dark brown, small (0.5-1.0 mm), irregularly circular and soli-
tary lesions on the abaxial leaflet surface. Initially these lesions are sub-
merged but later become raised and crust-like. Conidia of the fungus are
12-23 x 15.5-23.0 /-Lm in size, borne terminally, solitary, muriform, black,
and sometimes slightly rugose. Conidiophores are dark in colour and have
a globose apex and mayor may not be branched.
10.3.16 Choanephora leaf spot

Mukiibi (1975) reported Choanephora spp. causing leaf spots on ground-
nut in Uganda. The brown lesions originate at the leaflet margins and
spread to cover almost the entire leaflet. Abundant sporulation occurs on
Diseases of stems, roots and pods, caused by fungi 359
the adaxial and abaxialleaftet surfaces. Faint concentric circles are present
in the lesions.
10.4 DISEASES OF STEMS, ROOTS AND PODS, CAUSED BY

FUNGI
10.4.1 Stem rot

Stem rot caused by the fungus Sclerotium rolfsii Sacco is one of the most
widely recognized soilborne diseases of groundnut (Backman, 1984).
Other names for this disease include white mould, southern blight,
sclerotium rot, sclerotium blight, sclerotium wilt, root rot and foot rot
(Jackson and Bell, 1969). Yield losses vary between 25% and 80%
(Rodriguez-Kabana et al., 1975). The fungus attacks and kills the crown of
the plant, destroying entire plants and the pods they are producing.
First symptoms of the disease are usually the yellowing and wilting
of a branch or, if the main stem becomes infected, of the whole plant.
The leaves turn brown and sometimes are shed prematurely. Sheaths
of white mycelium can be seen at or near the soil line around affected
plant parts. Mycelial growth is rapid under favourable environ-
mental conditions and quickly spreads to other branches and plants.
Spherical sclerotia (0.5-2.0 mm in diameter) are produced abundantly
both on plant parts and on the soil surface. They are initially white
but later turn dark brown. Lesions produced on branches and pegs are
initially light brown, becoming dark brown as disease develops. Infected
pods usually rot and may occur on plants without visible symptoms
on the above-ground plant parts. S. rolfsii produces oxalic acid, a phyto-
toxin that gives a purple discoloration of seeds and is also responsible
for chlorosis and necrosis of foliage in the early stages of disease develop-
ment.
The fungus does not produce asexual spores. The basidial stage, Aethalia
rolfsii (Cruzi) Tu & Kimber, is rarely seen. S. rolfsii grows well on a wide
range of cultural media and is characterized by the presence of white
mycelium and hard, round, brown sclerotia.
S. rolfisi has a broad host range of more than 200 plant species (Aycock,
1966). Sclerotia are produced on many infected crops and weeds. Even in
the case of non-hosts, the debris from these plants may serve as an organic
food base for attack of subsequent crops. Warm, moist conditions favour
disease development. The fungus has a high demand for oxygen.
Therefore, the overwintering sclerotia are only activated when they occur
in the upper regions of the soil. Continuous cropping year after year may
cause inoculum build-up. The fungus is seedborne, in the form of my-
celium present both internally and externally, and may be carried through
seed to areas where the disease is not present. However, soilborne
360 Diseases
inoculum is the most important source of initial infection. Factors that tend
to increase or maintain soil moisture, such as a dense foliage canopy,
increase the disease severity (Backman et al., 1975).
Unfortunately, control of leaf spot diseases can directly increase the
severity of stem rot. Maintaining a complete canopy allows a moist sub-
canopy that is favourable to stem rot development. However, the results of
not controlling leaf spot diseases would be far worse (Backman et al.,
1975). Thus the application of chemicals to the soil is the most appropriate
approach to control this disease. It has been shown that application of
PCNB and carboxin could control the disease satisfactorily (Cooper, 1956;
Harrison, 1966), though at some locations responses with PCNB have been
erratic (Harrison, 1967; Indulkar and Grewal, 1970; Thompson, 1978).
Rodriguez-Kabana et al. (1979) obtained significant control by soil fumiga-
tion using dibromochloropropane.
Several cultural practices are effective for stem rot control. Deep
ploughing of soil after harvest to bury crop residues is commonly followed
in most of the groundnut-growing states in USA. One-year crop rotation
with maize (Zea mays) and grain sorghum (Sorghum bicolor) is effective in
preventing severe infestation. Rotations of 2-4 years are necessary if a
severe infestation has developed (Backman, 1984).
Porter et al. (1982) gave a detailed account of the biological control
of S. rollsii in groundnut. Trichoderma harzianum Rifai grown on gran-
ules containing molasses has been reported to be effective in controlling
the disease under field conditions (Backman and Rodriguez-Kabana,
1975).
10.4.2 Aspergillus species and aflatoxin

This subject is also considered in Chapter 13. In the early 1960s, mycotox-
ins of Aspergillus flavus Link were found in groundnut meal. Feed pre-
pared from this meal caused the death of 100000 turkeys in Great Britain,
and the disease was named Turkey X disease. Later a similar disease
caused deaths in ducklings in Kenya and Uganda. An extensive piece of
research indicated the presence of toxic Brazilian groundnuts used to feed
the birds at these places (Blount, 1961).
Groundnut pods and seeds are vulnerable to several soil fungi but
species of Aspergillus, Fusarium and Penicillium are the predominant
colonizers (McDonald, 1969; Mehan and McDonald, 1983) and produce
mycotoxins (Ciegler, 1978). Four mycotoxins - aflatoxin, citrinin, ochra-
toxin A and zearalenone - occur naturally in ground nuts and, of these,
aflatoxins produced by A. flavus and A. parasiticus Speare are the most
important and ubiquitous throughout the groundnut-growing areas of the
world (FAO/WHO/UNEP, 1977). Aflatoxins are toxic and even a small
amount of them in feed (10-20 ppb) can produce fatal liver cancer in young
animals and humans (Wogan, 1965; Diener et al., 1982).
A. flavus thrives in the soil as a saprophyte and invades groundnut seeds,
seedlings and pods during pre- and post -maturity stages in the field . Yellow
mould, a disease of ungerminated seeds and seedlings, and related afla-
toxin contaminations of groundnut seed can develop during harvest and in
storage (Dickens, 1977; Mehan and McDonald, 1983).
Seedling infection with A. flavus is characterized by necrotic lesions
on the emerging plumule and cotyledons. Lesions become covered with
masses of yellow-green spore heads of A. flavus group. Infection may
spread to the emerging radicle. Infected seedlings lack a secondary root
system, a condition known as 'aflaroot' (Chohan and Gupta, 1968; Aujla
et al., 1978). Infected seedlings sometimes die. The use of high-quality seed
treated with protectants greatly minimizes the severity of this disease.
A. flavus also attacks groundnut pods and seeds as they mature in the
soil before digging, especially when plants are drought-stressed. Yellow-
green mouldy colonies of Aspergillus develop on pods and seeds which are
overmature. Additional pods and seeds may become contaminated during
the curing process. The fungi invade maturing pods, seeds and cotyledons.
Hyphal colonies are apparent between the cotyledons. Aflatoxin contami-
nation during harvest, after harvest and in storage has had a tremendous
impact on the entire groundnut industry and on consumers. These aspects
of aflatoxin contamination have been adequately reviewed by several
workers (Mehan and McDonald, 1982; Mehan, 1988; Mehan et al., 1991),
who have also suggested possible control measures to minimize aflatoxin
contamination.
The taxonomy and morphology of the genus Aspergillus were described
by Raper and Fennell (1965). A. flavus and A. parasiticus produce hyphae
that are colourless, septate and branched. Conidiophores are 300-500 f-lm
long. A vesicle is borne at the end of each conidiophore. On the vesicle,
rows of strigmata bear chains of yellow-green to blue-green conidia (7-9 x
3.5-5.0 f-lm). Strigmata of A. flavus are borne in single or double series on
an elongate to subglobose vesicle (15-25 f-lm in diameter). The radiated
heads split with maturity. The strigmata of A. parasiticus are closely
packed in a single series on a flask- or pestle-shaped vesicle 20-35 f-lm in
diameter, and the heads are loosely radiate. Cultures of A. flavus produce
only aflatoxins Bl and B 2 , whereas cultures of A. parasiticus produce all
four aflatoxins (Diener et al., 1982).
The disease cycle and epidemiology of yellow mould and aflatoxin
contamination have been discussed in greater detail by Jackson and Bell
(1969) and Pettit (1984). A. flavus and A. parasiticus are saprophytic
inhabitants of many soils and survive in crop residues. Their ability to
cause plant disease is related to their competition with other soil micro-
organisms, the availability of susceptible plant tissues and occurrence of
favourable environmental conditions for infection and development of the
disease. In general, the A. flavus group of fungi is more competitive
in a declining soil moisture regime (Pettit et al., 1971) and when the
362 Diseases
atmospheric relative humidity remains ~90%. These fungi can grow over a
temperature range of 17--42 °C but the optimum temperature for aflatoxin
production is 25-35 0c.
An aflatoxin detection system has been developed in the United States
to prevent groundnut seed contaminated by aflatoxin from entering the
food chain of humans and animals (Dickens and Welty, 1969; Aibara and
Nobumitsu, 1977; Dickens, 1977). The FDA has set levels of aflatoxin for
the edible products trade.
Control of yellow mould and management of aflatoxin contamination of
groundnuts is achieved by preventing the A. flavus group from entering
groundnut tissues, by either destroying or diverting the contaminated seeds
and adopting improved crop husbandry (Dickens, 1977; Mehan, 1988).
Recommendations include:
• avoiding mechanical damage to the crop during cultivation, harvesting

and subsequent processing;
• harvesting at full maturity;
• drying the produce in the field as rapidly as possible;
• preventing re-wetting during or after drying;
• removal of damaged or mouldy pods;
• drying to a safe moisture level (8%) before placing in storage;
• storage at low temperature and humidity.
Such measures have been applied with considerable success by large-
scale groundnut farmers in the USA and other developed countries but are
neglected in developing countries (Mehan and McDonald, 1983).
Genetic resistance to seed invasion has been identified by several
workers (Mixon and Rogers, 1973; Bartz et al., 1978; Mehan et al., 1987)
but it depends upon the presence of an undamaged seed testa: any damage
to the testa greatly reduces the levels of resistance (Mehan, 1988).
10.4.3 Aspergillus crown rot

Aspergillus crown rot, a fungal disease caused by Aspergillus niger van
Tieghem, is found on groundnuts in all the major growing areas of the
world (Porter et al., 1982). The appearance of this disease on groundnut is
generally sporadic and unpredictable.
Although A. niger attacks groundnuts at all stages, seedlings and young
plants are particularly susceptible. Sudden loss of leaf turgidity and wilting
of plants are characteristics of crown rot. The crown of an infected plant
becomes swollen and the tissues become brittle and corky. Infected parts
are profusely covered with black masses of mycelium and conidia.
Additional symptoms of aspergillus crown rot are given by Garren and
Jackson (1973).
The fungus A. niger is widely distributed in soils throughout the world. It
colonizes and survives in organic matter. Growth and sporulation of the
fungus are usually favoured by warm and moist conditions, and under such
conditions it produces an abundance of large, black conidial heads that
reach 700-800 fLm in diameter. Soil type has not been correlated consist-
ently with prevalence of crown rot (Joffe and Lisker, 1970; Abdalla, 1974)
but the disease is more prevalent in soils low in organic matter (Morwood,
1953). A. niger is also capable of initiating disease through infected seeds
(Agnihotri and Goyal, 1971; Vaziri and Vaughan, 1976).
Most of the commonly grown cultivars are susceptible to the crown rot
fungus. However, bunch cultivars are usually less susceptible. Fungicidal
seed protect ants such as Thiram and Captan provide some control when
used under conditions and practices that favour rapid germination and
seedling emergence.
10.4.4 Blackhull
This disease is caused by the fungus Thielaviopsis basicola (Berk. and Br.)
Ferraris. In the United States the disease was first observed in 1963
(Mason, 1964) but there had been earlier reports from other countries
(Caccarone, 1949; Frezzi, 1960). More recently it has been reported from
South Africa (Prinsloo, 1980), where extensive hull discoloration, severe
pod and peg rots and up to 80% loss in pod yields were attributed to
blackhull disease.
Blackhull is characterized by scattered black dots on pod surfaces. As
individual lesions increase in size, they coalesce and form large blackened
areas. Large masses of chlamydospores embedded in the sclerenchymatous
hull tissues give rise to black discoloration of the hull.
Seed inside infected pods sometimes become discoloured and the causal
agent can often be isolated from these tissues. Tabachnik et al. (1979)
observed necrosis of the tap root and pegs.
Lucas (1965) described the morphological and physiological character-
istics of the causal organism. The fungus produces both endoconidia and
chlamydospores. Endoconidia are produced by specially modified conidio-
ph ores (4-6 x 9-14 fLm) composed of bulbous bases having a restricted
cylindric barrel.
T. basicola persists indefinitely in the soil as a saprophyte. It can
overwinter as chlamydospores in the endocarp of the fruit (Mason, 1964).
Resting chlamydospores germinate and produce mycelium which grows
along the young pod surface and penetrates the endocarp. Later, chlamy-
dospores are produced abundantly from the mycelium in a characteristic
black granular mass which may fill the intercellular spaces of the mesocarp.
Hsi (1965) found no constant relationship between seed from infected pods
and subsequent disease incidence.
Crop rotation with grain sorghum controls blackhull disease (Hsi, 1978),
and it can be suppressed if benomyl and thiophanate-methyl are applied
364 Diseases
into the soil (Hsi and Ortiz, 1980). However, no commercial groundnut
cultivars are resistant to this disease.
10.4.5 Botrytis blight

Botrytis blight of groundnuts is caused by Botrytis cinerea Pers. ex. Fries.
It was first reported in Japan (Suematu, 1924) and has been observed in
most of the groundnut-producing countries worldwide (Porter, 1984c).
Occasional outbreaks do substantially reduce yields. The disease is also
called 'grey mould' and in South Africa it is known as 'botrytis shoot
disease' (Dyer, 1951).
All mechanically wounded groundnut plant parts are invaded by this
soilborne fungus. Tissues weakened by other fungi (Rothwell, 1962) or
injured by frost are vulnerable to colonization by B. cinerea. Under
favourable environmental conditions, the fungus colonizes plant tissues
rapidly and moves from above-ground plant parts into underground parts.
H produces abundant conidia and sclerotia on infected tissue. Conidia are
hyaline, ellipsoid to ovoid, single-celled, measuring 9-12 x 6.5-10 /-Lm.
Conidiophores are usually unbranched septate and 11-23/-Lm thick.
The ascomycetous stage of the fungus is Botryotinia fuckeliana (de Bary)
Whetzel.
Conidia are frequently disseminated by air currents. Low temperatures
(~20 °C) accompanied by heavy dews or excessive rainfall favour infection
and disease development. The fungus overwinters in the sclerotial form in
the soil. Although apothecia have been reported, the primary source of
inoculum appears to be mycelium originating from germinating sclerotia or
omnipresent conidia.
Fungicides such as benomyl, chlorothalonil and dicloran offer some
control when sprayed on groundnut foliage. Early-maturing groundnut
cultivars may escape frost damage and thus reduce disease severity in some
countries.
10.4.6 Charcoal rot

Charcoal rot, caused by a common soilborne fungus, Macrophomina pha-
seolina (Tassi) Goid., is widely distributed throughout the world's
groundnut-producing countries. The fungus has a wide host range and in
groundnuts it causes seed and seedling rots, wilt, root and stem rot, leaf
spots, and rotting of developing pods and seeds. The term 'charcoal rot' is
used to describe the damage done to roots and stems of seedlings and older
plants. It is a disease of minor importance but can greatly reduce seedling
stands (Hoffmaster et al., 1943; Mathur et al., 1967; Shanmugam and
Govindaswamy, 1973a).
Groundnut seedlings are usually attacked by the fungus at the soil line,
and water-soaked lesions appear on the hypocotyl. Later, infected hypo-
cotyls are girdled and the seedlings die. Similar symptoms are observed on
the older plants at the soil line, although all plant parts at all stages of
growth are vulnerable to attack. Stem and root lesions appear water-
soaked at first, but infected tissues later become a dull light brown. The
infection extends downwards into the taproot and up into the stem and
branches. Where lesions girdle the stem, the plant wilts and the fungus
rapidly colonizes the branches. Jackson and Bell (1969) published an
excellent summary of symptoms caused by M. phaseolina.
Pycnidia of M. phaseolina are membranous to sub-carbonaceous, first
immersed and then at least partially erumpent, globose or flattened globose
with inconspicuous truncate ostioles, 100-200 J.Lm in diameter. Walls are
composed of several layers of dark thin-walled angular cells, 9 J.Lm in
diameter, and are lined with a hyaline layer two or three cells thick, bearing
simple rod-shaped conidiophores, 10-15 J.Lm long. Conidia (14-33 x 6-12
J.Lm) are single-celled, hyaline and elliptic or oval. The sterile mycelial phase
of the fungus is commonly known as Rhizoctonia bataticola (Taub.) But!.
Charcoal rot is both seedborne (Subrahmanyam and Rao, 1977; Mridha
and Fakir, 1978) and soilborne (Garrett, 1956). Mycelium in seeds and
mycelium and sclerotia in plant debris are the primary sources of inoculum.
The sclerotia can remain viable in dry soil for many years (Smith, 1966),
but soon lose viability in very wet soils (Shokes et al., 1977). The disease
can readily be disseminated by infected groundnut seeds and pods. High
soil temperatures, low soil osmotic potential and reduced plant vigour
favour the development of charcoal rot (Odvody and Dunkle, 1979).
Crop rotation is generally ineffective in reducing the soil inoculum
because the fungus can grow saprophytically and has a wide host range.
Adequate fertilization and soil water to ensure good crop growth reduce
the chances of infection and charcoal rot development. Soil drenching with
PCNB has given some control (Shanmugam and Govindaswamy, 1973b).
No immune or highly resistant genotypes have been identified.
10.4.7 Diplodia collar rot

Diplodia collar rot, caused by Diplodia gossypina (Cke.) McGuire and
Cooper, occurs sporadically throughout groundnut-growing countries of
the world but rarely causes economically important losses. The disease is
also known as 'diplodia blight' and 'collar rot'.
Groundnut seedlings and older plants are attacked at or near soil level
and the fungus quickly invades the stem. The first obvious symptom of the
disease is wilting of a lateral branch or the entire plant. Wilt develops
rapidly and plants generally die within a few days. Lesions that develop on
the above-ground tissues are characterized by elongated necrotic areas
having light brown centres and dark brown margins. Infected roots become
slate grey to black and shred easily. Pycnidia on the infected host tissue
366 Diseases
appear as small black dots. The fungus is capable of causing concealed
damage to the seeds (Porter, 1984b).
Morphology and physiology of the causal organism are described by
Jackson and Bell (1969). Although several species of Diplodia have been
isolated from infected groundnut parts, D. gossypina is probably the most
common and the causal agent of collar rot disease. Simple or compound
pycnidia occur either singly or in groups over the necrotic tissues. Pycnidia
range up to 400 J.Lm in diameter. Conidia are borne on short conidiophores;
mature conidia (17-34 x 10-18 J.Lm) are elliptic with one septation. They
are brown with longitudinal striations.
D. gossypina survives as mycelium and mature conidia in the soil and in
crop residues for a long time. Upon germination, mature conidia may
initiate primary infection. Groundnut plants predisposed by heat injury are
more vulnerable but predisposition is not a prerequisite for infection. Once
the infection site is established, the fungus moves quickly into the adjacent
healthy tissue. Under field conditions hot dry weather favours disease
development.
Crop rotations with non-host crops such as cotton or soybean can reduce
disease incidence. Resistance to D. gossypina is not found in commercial
cultivars but some breeding lines that possess high levels of tolerance to the
disease are available in the USA (Porter and Hammons, 1975; Porter,
1984b). Predisposition of groundnuts to D. gossypina by heat injury can be
minimized by controlling foliage diseases so that a dense foliage canopy
shade is maintained throughout the growing season.
10.4.8 Pythium diseases

Several species of Pythium cause pod breaking (pod rot), pre-emergence
and post-emergence damping-off, vascular wilt and root rot diseases of
groundnuts throughout the world. All Pythium species are cosmopolitan in
soils and attack a wide range of crop species.
Pod breakdown or pod rot (Garren, 1966) is widespread in groundnut-
growing areas of the world and often causes economic losses that are
difficult to define, because infection by Pythium spp. and other related
fungi usually does not produce well-defined above-ground symptoms.
P. myriotylum Drechs. is the major pod-rotting pathogen in certain
groundnut-growing areas in the USA. However, several other species of
Pythium, Rhizoctonia solani and Fusarium solani can cause pod rot either
singly or in combination. The synergistic effects of P. myriotylum and
F. solani in a pod breakdown complex are known in Israel (Frank, 1972).
In Israel, Pythium spp. are thought to precede Fusarium spp. (Frank,
1968), while in the United States Fusarium spp. usually precede Pythium
spp. (Garren, 1966). P. myriotylum attacks groundnuts at all stages of
growth (Frezzi, 1956).
Pod breakdown symptoms caused by the above-mentioned fungi, either
singly or in combination, are often indistinguishable. Both mature and
immature pods may be attacked. Infected pods exhibit degrees of dis-
coloration from apparent rusting to browning. In wet soil, pegs and pods
infected by P. myriotylum become water-soaked. They later become dark
brown to black and rot quickly (Jackson and Bell, 1969). F. solani and R.
solani produce distinct lesions under dry soil conditions. Pod rot caused by
R solani progresses much more slowly than that caused by P. myriotylum
(Garren, 1970; Garcia and Mitchell, 1975a).
Occasionally groundnut plants without apparent pythium root rot symp-
toms wilt suddenly (Porter, 1970). Shortly after the appearance of wilt
symptoms, leaflets become chlorotic or light green. This condition of the
diseased plants is commonly referred to as 'vascular wilt'. The plants have
some evidence of root deterioration; vascular tissues of the tap root are
dark brown. Fibrous roots are also vulnerable to P. myriotylum and
infected roots become dark brown to black (Jackson and Bell, 1969).
P. myriotylum was described by Drechsler (1930), and a study of the
genus Pythium as plant pathogens was made by Hendrix and Campbell
(1973). Pythium spp. are easy to isolate from infected tissues and they are
characterized by the presence of coenocytic mycelium, from which develop
asexual reproductive structures (sporangia) that differ in size and shape
between species. The sporangia of P. myriotylum may be terminal or
intercalary and germinate by germ tube or by producing zoospores.
Zoospores and sporangia are short-lived. The primary survival structures
of P. myriotylum in soil are oospores (sexual spores), which are 12-37 f.Lm
in diameter and do not fill oogonia. Oospores have a single reserve
globule, walls up to 2 f.Lm thick and pale golden content.
Pythium spp. are natural inhabitants of the soil and can survive indefi-
nitely as saprophytes. P. myriotylum has a wide host range that includes
cereal crops used in rotation with groundnuts (McCarter and Littrell,
1968). Mycelia of P. myriotylum, produced by zoospores or germination of
oospores, form appressoria and penetrate epidermal cells of groundnut
pods directly (Jones, 1975). Penetration occurred in 2 hours at 30-34 DC
but there was no penetration below 22 DC. The optimum temperature for
mycelial growth of P. myriotylum is 35 DC.
A significant positive correlation between increasing soil moisture and
frequency of pythium pod rot has been demonstrated in Israel (Frank,
1974). Frequent irrigation of sandy soils increased the severity of pod rot
caused by a combination of P. myriotylum and F. solani, whereas less
frequent and heavier irrigations reduced its severity. The soil fauna, in-
cluding mites and springtails (Shew and Beute, 1979), nematodes (Garcia
and Mitchell, 1975a) and southern corn root-worm (Porter and Smith,
1974), enhances pod breakdown disease caused by P. myriotylum.
Moderate levels of resistance to pod rot caused by P. myriotylum and
high yield potentials have been reported (Porter et ai, 1975).
Since more than one organism is involved in pod breakdown, wide-
368 Diseases
spectrum fungicides or combinations of fungicides are needed to control
pod rot. Use of effective nematicides can be important, since some types of
nematodes intensify pod rot in certain locations (Garcia and Mitchell,
1975b). Pod breakdown caused by P. myriotylum can be significantly
suppressed by the application of high rates of gypsum (Garren, 1964;
Hallock and Garren, 1968; Boswell and Thames, 1976). Control of
pythium diseases of groundnuts through crop rotation and field manage-
ment have been difficult.
10.4.9 Cylindrocladium black rot

Cylindrocladium root, peg and pod rot caused by the fungus
Cylindrocladium crotalariae (Loos) Bell and Sobers (perfect stage,
Calonectria crotalariae (Loos) Bell and Sobers) is reported to occur in the
United States (Bell and Sobers, 1966), Japan (Misonou, 1973), Australia
(Jackson and Bell, 1969) and India (Sharma et al., 1978). The disease has
the capacity to inflict heavy yield losses (Garren et al., 1972; Lewis et al.
1977).
The disease was reviewed in 1980 (Beute, 1980). The first symptoms of
cylindrocladium black rot (CBR) are wilting and chlorosis (yellowing) of
infected leaves and stems. Usually the central stem dies. As the disease
progresses, the lateral branches also die. Occasionally the whole plant
becomes chlorotic and stunted. All underground plant parts may develop
CBR symptoms. Hypocotyl, roots and pods become black and necrotic. A
diagnostic sign of CBR is the occurrence of small, reddish-orange perithe-
cia of the pathogen in dense clusters on stems, pegs and pods. These
structures form only on tissues within a few millimetres above or below the
soil surface during periods of wetness and high humidity. If these symp-
toms are lacking, a positive diagnosis of CBR requires a laboratory assay.
When CBR is seen for the first time in a field, the infested area develops in
one or more localized spots, which sometimes measure 10-30 m in
diameter.
The fungus grows well on potato-dextrose agar, producing light yellow
to white web-like aerial mycelium and a burnt orange to dark brown
submerged growth. Morphology and taxonomy of both sexual and asexual
stages have been described by Bell and Sobers (1966).
Microsclerotia of C. crotalariae are produced abundantly in roots and
Rhizobium nodules. As the infected tissue decomposes, microsclerotia are
released into soil and disseminated by tillage equipment. They overwinter
in soil, and soil temperature during winter months is a primary determinant
of longevity. Their population declines sharply in frozen soil (Phipps and
Beute, 1977) but they can survive for several years in the soil without a host
crop. Development and ejection of ascospores from perithecia of C. crota-
lariae was optimum at 25°C (Rowe and Beute, 1975). Ascospores are
extremely susceptible to desiccation and thus play only a minor role in
disease spread. Wind and water can contribute to long-range spread of the
pathogen.
The development and release of CBR-resistant cultivars of groundnut
appears to be one of the most promising strategies for CBR control. In
general, spanish cultivars are most resistant, valencia are the least resistant
and virginia cultivars are intermediate.
Crop rotations that include non-legumes such as corn, sorghum or forage
grasses minimize the development of new destructive races of the patho-
gen. Sanitation can be used to minimize infection by C. crotalariae.
Movement of the pathogen from field to field can be reduced by cleaning
tillage equipment (Krigsvold et al., 1977) and by leaving groundnut debris
on the soil surface for maximum winter kill of microsclerotia.
10.4.10 Rhizoctonia diseases

Groundnut diseases caused by Rhizoctonia solani Kuhn (teleomorphic
stage, Thanatephorus cucumeris [Frank] Donk) are commonly known as
damping off, root and stem rot, pod breakdown (pod rot) and foliage
blight. They occur throughout the world (Porter et al., 1982). The econ-
omic importance of these diseases is difficult to assess because they usually
occur simultaneously with other soilborne diseases and foliar pathogens.
However, diseases caused by R. solani have become increasingly important
with the greater use of irrigation and cultural practices that produce dense
canopy growth.
Groundnut seeds are frequently infected by R. solani before or shortly
after germination. It may cause damping off of the seedling or dry sunken
cankers on the hypocotyl (sore skin) and taproot. Later in the growing
season the fungus forms infection cushions on the hypocotyl, penetrates
the epidermal and cortical cells and causes the tissues to collapse.
Roots infected by R. solani develop small sunken lesions of a light to mid
brown colour. The root cortex of young seedlings is decayed and necrosis
frequently envelops the entire root system, extending up the hypocotyl and
killing the plant.
Branches of the mature plant that are in contact with the soil are freely
colonized by R. solani. Lesions, which are sunken and circular at the site of
infection, elongate and girdle the stem. The infected branch wilts and dies.
With abundant moisture, the fungus often grows to the top of the foliar
canopy, destroying leaves and stems. Leaf lesions are light to dark brown
and have distinct zonate patterns.
R. solani frequently infects the tips of pegs just below the soil surface and
destroys the pod-forming region. If soil moisture and temperature are
favourable, the fungus may grow up the pegs and produce sunken and
elongated lesions. Pegs are girdled and tissue becomes shredded with
exposed vascular bundles.
Pods may be infected by R. solani at any stage of development. Infected
370 Diseases
pods have dull coloured (light to dark brown) sunken lesions in contrast to
those infected with P. myriotylum, which have lesions that appear black
and greasy. This phase of the disease is frequently referred as pod break-
down (pod rot).
R. solani is usually found without a distinct spore form and thus is put in
the group of fungi known as Mycelia sterilia (Ainsworth, 1963). It produces
many sclerotia in host tissue. Sclerotia, which are aggregates of thick-
walled hyphae, can persist in the soil in the absence of the host crop (Blair,
1943; Papavizas and Davey, 1960). Sclerotia germinate when stimulated by
exudates from a susceptible host or by the addition of organic matter to the
soil. If the soil contains adequate organic matter, the fungus can grow
saprophytically. Since many crops and weeds are susceptible to R. solani
and organic debris is periodically added to the soil, long-term survival of
the fungus may occur in many soils. R. solani is a common contaminant of
groundnut seed and may initiate seedling diseases. Movement of soil or
infested plant residue also facilitate the spread of the fungus (Garren and
Jackson, 1973).
Resistance to seedling and pod diseases caused by R. solani is available
in wild groundnuts but not in commercial cultivars. Chemical seed protec-
tant may provide some control of seedborne infection (Jackson, 1963). The
systemic fungicides benomyl and carboxin (Abu-Arkoub, 1973), TCMTB
(Apdou and Khadr, 1974) and a combination product of thiram, picloram
and carboxin provide moderate to good control of seed and seedling
diseases caused by R. solani. Cultural practices also aid in controlling
groundnut diseases (Garren and Jackson, 1973).
10.4.11 Sclerotinia blight

Sclerotinia blight was first observed on groundnut plants in Argentina in
1922. It is now present in most groundnut-producing countries of the world
(Porter, 1980b). Yield losses of 10% are common, and in fields showing
severe symptoms of disease pod losses often exceed 50% of the expected
yield (Porter, 1980b).
Sclerotinia minor Jagger is the causal agent but S. sclerotiorum (Lib) de
Bary has also been found associated with this disease. The first symptoms
are lighter green to yellowing of foliage of plant tops and loss of turgidity
(flagging or wilting). Examination of the lower canopy of the infected plant
early in the morning reveals the presence of fluffy cottony mycelia of the
fungus on the main stem or lateral branches and other affected parts.
Initial loci are characterized by small, light green, water-soaked lesions,
which later appear sunken and become elongated and light tan. Older
lesions are dark brown with a distinct zonation between infected and
healthy tissues.
Foliage of the infected branches becomes chlorotic, turns dark brown,
and withers. Once the stem is girdled, the branch dies. These symptoms
result in the blighted appearance for which the disease is named. Shredding
of infected stems, branches and pegs is a characteristic sign of this disease;
another prominent characteristic is numerous sclerotia on and in infected
plant tissues.
In nature, S. sclerotiorum is always found in conjunction with S. minor.
In artificial inoculations, either species can produce blight lesions.
Apothecia of S. minor are rarely observed in soils during growing season.
However, apothecia are generally observed on crop debris; they are pale
orange to white and have a funnel-shaped or flat top. Kohn (1979) used
several characteristics to resolve the taxonomic position of the genus
Sclerotinia. Additional taxonomical and morphological characteristics to
support the separation of Sclerotinia species are provided by Willetts and
Wong (1980).
Sclerotia of the fungus can persist in field soils for 4-5 years in the
absence of groundnuts. One sclerotium in 100 g of soil is reported to be
sufficient to initiate the disease (Porter, 1980b). This may result in severe
disease in groundnut when environmental conditions are conducive to its
development. Infection with S. minor is favoured by cool temperatures
(18-20 0q, moist soil, high rainfall and high relative humidity (95-100%)
in the lower crop canopy (Porter, 1980b).
Tillage practices such as ploughing distribute sclerotia throughout the
field. Mechanically injured groundnut foliage is very susceptible to coloniz-
ation by S. minor (Porter and Powell, 1978). Plants injured by tractor tyres
during pesticide application were attacked at twice the frequency of nonin-
jured plants.
Effective control measures are not yet available (Porter, 1984a). The use
of some systemic fungicides, soil fumigants and biocides can aid in reducing
losses due to disease (Beute et at., 1975; Porter, 1977; Porter, 1980a) but
systemic fungicides such as chlorothalonil or captafol, when applied to
control leaf spot fungi, increase the severity of sclerotinia blight (Porter
1980a). Cultural practices such as crop rotation with a non-leguminous
crop (corn or sorghum), and avoiding excessive irrigation during the
growing season when temperatures are cool, help to reduce the soilborne
inoculum of the pathogen.
10.4.12 Verticillium wilt

A vascular wilt of groundnuts caused by a soilborne fungus Verticillium
dahliae Kleb. was reported from Asia (Golovin, 1937; Morwood, 1953),
USA (Smith, 1960) and Australia (Purss, 1961; Purss, 1962). The disease
occurs in other groundnut-growing countries but is not considered of
economic importance worldwide. However, it can cause yield losses as
high as 50% (Smith, 1960; Purss, 1961).
The first symptoms of verticillium wilt appear at flowering time but older
plants are also infected. Symptoms consist of marginal chlorosis of leaflets,
372 Diseases
loss of leaf turgidity and curling of leaves. Leaflets of the infected plants
usually turn dull green. The root systems of infected plants appear normal
except for a brown-to-black vascular discoloration. Yellowing, leaf necro-
sis, wilting, defoliation, stunting and eventual dehydration of infected
plants become more pertinent as the disease progresses.
Though V. dahliae is considered as the primary causal agent of verticil-
lium wilt, V. albo-atrum Reinke and Berth has also been found to cause
wilt in groundnuts (Smith, 1960). Both the species have a broad host range
and are widely distributed. The taxonomic basis for separation of the
species has been questioned (Jackson and Bell, 1969).
V. dahliae can survive in the soil for long periods by forming black
microsclerotia which can resist adverse environmental conditions. These
microsclerotia remain dormant until groundnut root exudates stimulate
them to germinate. The fungus initially invades plants through their root
system, then develops rapidly and systemically, spreading throughout the
vascular system and all plant parts. The pathogen spreads by movement of
farm machinery from infected to non-infected fields and also by wind and
water-borne movement of infected soil and plant tissue. Another possible
mode of spread is through infected seeds.
Populations of wilt pathogens can be suppressed by rotation with non-
host crops such as grain sorghum and alfalfa (Hsi, 1967). Removing and
burning of crop residues helps in reducing the inoculum density of the
pathogen in field soils. Use of clean verticillium-free seed is also recom-
mended. Chemical control of the disease has not been satisfactory in the
USA but fungus was effectively controlled in sandy soils in Israel with
metma-sodium applied through sprinkler irrigation (Krikun and Frank,
1981). Adequate levels of resistance to this disease are not available (Khan
et al., 1972).
10.5 DISEASES CAUSED BY NEMATODES
Diseases caused by plant-parasitic nematodes are among the least under-

stood problems currently confronting agriculture. Yield losses from nema-
tode diseases are generally due to their feeding on the roots, which
weakens the plants and reduces their vigour. Groundnut is one of the few
crops in which nematodes reduce crop yield directly by damaging pegs,
pods and seeds, and indirectly by feeding on roots and weakening the
plants. On a worldwide basis, annual groundnut yield losses caused by
nematodes are estimated at approximately 12% , while monetary losses are
estimated at just over one billion US dollars (Sasser and Freckman, 1987).
Although more than 90 species of plant-parasitic nematodes have been
reported in association with groundnut (Sharma and McDonald, 1990a),
only a few species are known to cause economically important diseases
(Subrahmanyam et al., 1990). Species of Meloidogyne, Pratylenchus,
Diseases caused by nematodes 373
Aphasmatylenchus, Scutelionema, Belonolaimus, Criconemelia, Tylencho-
rhynchus, Aphelenchoides and Ditylenchus can cause appreciable damage
to groundnut and are considered to be economically important in several
groundnut-producing regions.
10.5.1 Pod and root-knot disease

This disease, caused by four species of Meloidogyne (M. arena ria (Neal)
Chitwood, M. hapla Chitwood, M. incognita Kofoid and White, and
M. javanica (Treub) Chitwood), is the most important nematode disease of
groundnut. It is widespread over the groundnut-producing regions of the
world. M. arenaria and M. javanica are more common in the warmer
regions, and M. hapla in the cooler regions. M. incognita occurs in the
Mediterranean region (McDonald and Raheja, 1980).
M. arenaria is the most widespread and serious of the groundnut root-
knot nematodes. In Senegal, it does not successfully parasitize groundnut;
juveniles penetrate the roots and subsequently die (Netscher, 1975).
However, groundnuts sown in soils heavily infested with this nematode
have crop establishment problems as numerous juveniles penetrate the
roots, causing browning and necrosis of tissues and in some cases leading to
death of the young plants.
Root-knot damage to groundnut is often not suspected until roots are
examined and the typical galls are observed on roots and pods. Galling can
occur on all underground parts including the pods, which appear warty.
Nematode infection disrupts the vascular system, resulting in devitalized
roots, retarded growth, yellowing of foliage, stunting and wilting of plants.
Gall size varies depending on the nematode species involved. M. hapla
produces small galls and extensive root proliferation - roots often form a
dense mat when infection is severe. Production of roots just above the gall
is generally associated with M. hapla infection. Galls produced by
M. arenaria and M. javanica infection are very similar to one another.
Early infection of pegs prevents full development of the fruits so that
diseased plants have many necrotic pegs and few mature pods.
Damage to groundnut roots by M. arenaria at an early stage in plant
development impairs growth of the main taproot. As the root system is
small at this time, this affects the plant growth more severely than damage
which occurs later in the season when roots are well developed and are
capable of sustaining the nematode population without serious effects on
plant growth. Loss in yield can occur even in soils harbouring as few as 0.5
juveniles/cm3 of soil.
Nematode infection reduces the number and efficiency of Rhizobium
nodules. M. arenaria infection increases the incidence of damping-off
disease of groundnut caused by Pythium myriotylum Dreschler (Garcia
and Mitchell, 1975b). Inoculation of groundnuts with M. arenaria and
Fusarium solani (Mart.) Sacco together causes earlier appearance of wilting
374 Diseases
of groundnut than when F. solani is inoculated alone (Patel et al., 1985).
The nematode also enhances development of cylindrocladium black rot
[CO crotalariae (Loos) Bell and Sobers] on the resistant variety NC 3033.
M. hapla similarly enhances the development of cylindrocladium black rot
(Diomonde and Beute, 1981). Soil population densities of M. arenaria at
more than 0.01 individual/cm 3 in the USA and 1.0 individualslcm 3 in India
at sowing can cause appreciable damage (Sharma and McDonald, 1990b).
Production losses due to M. arenaria in major ground nut-producing
states of the USA range from 0.5% to 5.4%, and for M. hapla range from
0.3% to 4.7% (Anon., 1987). Estimates of economic losses caused by pod
and root-knot are not available from many countries but the disease is
considered important in Australia, the People's Republic of China, Egypt
and India (Sharma and McDonald, 1990b).
Farmers in Egypt and in the USA use nematicides to control pod and
root-knot disease. Aldicarb, carbofuran, ox amyl and phenamiphos at the
rate of 1.0-3.4 kg a.i.lha are generally used (Rodriguez-Kabana and King,
1979; Rodriguez-Kabana and King, 1985; Rodriguez-Kabana et al., 1981).
Seed treatments with 3% aldicarb sulfone and 6% carbofuran reduce the
damage by M. arenaria (Patel et al., 1986). Formulations of fumigant
nematicides are either not available in developing countries or their use is
restricted.
Rotations with non-hosts or poor hosts can be effective in reducing the
damage to groundnut. The root-knot species have a wide host range but
monocotyledons are, in general, non-hosts or poor hosts of the species that
attack groundnut, so that maize, sorghum, pearl millet, and rice are good
crops to rotate with groundnut. Sesamum indicum L., Ricinus communis
L., Aeschynomene indica L., Cassia fasiculata Michx., Indigofera hirsuta
L., and Paspalum notatum Flugge are also good crops to include in
rotations for reducing M. arenaria populations (Rodriguez-Kabana and
Morgan-Jones, 1987; Sasser and Kirby, 1979). Appropriate rotations
should reduce population densities below the damage threshold. Breaks of
three years or more between susceptible crops in the rotation should give
good control of root-knot disease. Care should be taken to remove weed
hosts of the root-knot species during this break period.
The ideal method for controlling root-knot nematodes on groundnut
would be to use resistant cultivars; however, only limited information is
available on this subject. In most countries where the disease is important,
no efforts are being made to identify resistance. Resistances in wild
Arachis spp. have been identified to M. arenaria and M. hapla (Baltens-
perger et al., 1986; Castillo et al., 1973; Holbrook and Noe 1990; Nelson et
al., 1988). Eleven groundnut lines (NC 8C, NC 6, NC 343, NC 3033, NC
17921 x NC 18016, Tainan 9, CES 103, CS 20, NC Ac 17090, NC 18230 x
NC 2 and NC 18231 x NC 2) have been reported to be resistant to M.
arenaria and M. hapla (Anon., 1985). Grewal et al. (1986) reported JSP 1,
ICGS 2, NC 4X and CGC 4 to be resistant to M. arenaria, and Sakhuja and
Sethi (1985) found four genotypes resistant to M. javanica. Resistance of
these varieties needs to be tested under field conditions.
10.5.2 Pod and root-lesion disease

This is an important nematode disease of groundnut in Australia, Brazil,
Thailand and Zimbabwe. Species of Pratylenchus, Tylenchorhynchus,
Criconemella, and Belonolaimus cause lesions on pods and/or roots, but
Pratylenchus is the most important. Steiner (1949) first reported
Pratylenchus brachyurus (Godfrey) Filipjev and Sch. Stekh. on groundnut
in the USA, and this species is an economic deterrent to groundnut
production in Texas (Smith et al., 1978). The nematode was recently
reported in India from one of the major groundnut-producing areas of
Andhra Pradesh (Varaprasad and Sharma, 1990) but its distribution in
other Indian states is not known.
Belonolaimus longicaudatus Rau occurs in sandy soils along the Atlantic
coastal plain of the USA from New Jersey to Florida and westward to
Texas and Arkansas (Minton and Baujard, 1990). This nematode has not
been found outside the USA.
Criconemella ornata (Raski) de Grisse and Loof causes lesions on pods
and roots of groundnuts in the USA and has been associated with ground-
nuts in the African countries of Burkina Faso, Egypt and the Gambia, and
in India (Germani and Dhery, 1973; Ibrahim and El-Saedy, 1976; Sharma,
1988). Tylenchoryhnchus brevilineatus Williams damages groundnuts in
the Nellore and Chittoor districts of Andhra Pradesh, India (Reddy et al.,
1984). Distribution of this nematode-caused disease is apparently limited
to Andhra Pradesh.
P. brachyurus feeds on cortical cells. The enzymes produced by the
nematode hydrolyse storage compounds to toxic products which cause
lesions. These start as small brown flecks where the nematode enters, and
increase in size with time. Once a lesion appears, fungi and bacteria attack
the dead tissue and cause rotting. Pratylenchus spp. are migratory endo-
parasites and are carried in roots and pegs, and in the shells of mature
pods. Several hundred nematodes may be present in a single lesion. It is
strongly suspected that P. brachyurus increases the severity of diseases
caused by Sclerotium rolfsii Sacc., Pythium sp. and Aspergillus fiavus Link:
Fr. (Jackson and Minton, 1968; Sharma and McDonald, 1990b).
B. longicaudatus is an ectoparasite and feeds on root tips as well as on
young pegs and pods. Small necrotic lesions may be observed on the roots,
pegs and pods (Owens, 1951). The nematode-infected plants are severely
stunted and chlorotic (Minton and Baujard, 1990). Heavy infestations
cause stubby lateral roots. Roots, pegs and pods attacked by C. ornata are
severely discoloured with brown necrotic lesions and many lateral root
primordia are killed (Minton and Bell, 1969).
T. brevilineatus infection causes brownish black discoloration of the pod
376 Diseases
surface. Small brownish-yellow lesions appear on pegs and young develop-
ing pods. Discoloration on roots is less severe than on pods (Reddy et al.,
1984). Above-ground symptoms of T. brevilineatus are often deceptive;
foliage is dark green and this distinguishes the disease from that caused by
C. ornata which leads to yellowing of foliage on diseased plants.
Damage thresholds for P. brachyurus are not well established.
Application of nematicides in infested soils in the USA results in significant
increase in yield (Boswell, 1968; Minton and Morgan, 1974). Although
most infestations cause only moderate to low damage, the nematode is
considered important in the USA because of its widespread distribution in
groundnut soils (Ingram and Rodriguez-Kabana, 1980) and production
losses due to Pratylenchus spp. range from 0.1% to 2.0% (Minton and
Baujard, 1990). No information has been obtained from other countries.
Sasser et al. (1960) obtained yield increases of more than 100% with
nematicide application in fields infested with B. longicaudatus. Economic
losses in the USA due to this nematode range from 0.25% to 0.5% (Anon.,
1987). Losses due to C. ornata are not well established and damage in the
field often goes undetected. A population density of less than one C.
ornata individual/cm 3 soil can cause significant loss in yield (Barker et al.,
1982) and one T. brevilineatus individual/cm 3 of soil causes significant loss
in yield.
Nematicides that control the pod and root-knot diseases also control the
pod and root-lesion diseases but chemical control is not commonly prac-
tised. For B. longicaudatus, however, nematicide application is a major
means of control (Minton and Baujard, 1990). Crop rotations are not
entirely effective because of the wide host ranges of the nematodes.
Fallowing for more than 6 weeks greatly reduces P. brachyurus popu-
lations (Brodie and Murphy, 1975). There are no commercial cultivars with
resistance to P. brachyurus, C. ornata or T. brevilineatus. Three plant
introductions to the USA - PI 1290606, PI 1295233 and PI 1365553 - are
resistant to P. brachyurus (Smith et al., 1978; Starr, 1984) and preliminary
field screening work indicates the availability of 23 sources of resistance to
T. brevilineatus in groundnut genotypes (Siva Rao et al., 1986; Mehan and
Reddy, 1987).
10.5.3 Groundnut chlorosis

This disease, caused by Aphasmatylenchus straturatus Germani, is confined
to south-west Burkina Faso where it has infested approximately 2600 ha of
groundnut crops (Minton and Baujard, 1990). The nematode causes inter-
veinal chlorosis and stunting of plants. Chlorotic plants have reduced root
systems and poor nodulation (Germani and Dhery, 1973).
Disease symptoms occur in fields with less than one A. straturatus
individual/cm 3 soil. Yield reductions due to groundnut chlorosis range
from 30% to 70% .
Application of dibromochloropropane (DBCP) at sowing gives satisfac-
tory control of the disease (Dhery et al., 1975). The nematode attacks
many economically important crops in Burkina Faso (Germani and Dhery,
1973). There is no information on crop rotations likely to reduce disease
severity, nor are there reports of varietal resistance.
10.5.4 Yellow patch disease/crop growth variability

This is a problem in the Sahelian Zone of West Africa and two species of
Scutellonema have been implicated as major biotic factors (Germani et al.,
1985; Sharma et al., 1990). Scutellonema cavenessi Sher in Senegal and
S. clathricaudatum Whitehead in Niger cause serious damage to groundnut
crops and the symptoms in each country are similar. These species are also
reported from other countries in West Africa where they are suspected of
causing damage to groundnuts (Sharma, 1989). Foliage of groundnut
plants grown in soil infested with the Scutellonema spp. is chlorotic with
reduced leaf size and canopy development. Lateral roots are also reduced
in size. The poorly growing plants appear in randomly distributed patches
which persist and increase in size during the subsequent growing season.
The extent of crop loss in Senegal and Niger has not been fully evalu-
ated. Application of nematicides to the nematode-infested fields in these
two countries have increased pod and haulm yields by more than 100%
(Germani et al., 1985). An initial population density of more than one
S. clathricaudatum individual/cm 3 soil causes stunting of foliage. The
nematode-caused damage appears to be aggravated by low soil pH and
aluminium toxicity in the soil (Sharma, 1989). Applications of DBCP at
151/ha and carbofuran at 6 kg a.i.lha reduce the nematode populations in
soil and increase crop growth and yield (Germani and Gautreau, 1976;
Sharma, 1989). The are no known groundnut cultivars resistant to
S. cavenessi, but screening of cultivars for resistance or tolerance to
S. clathricaudatum has shown that some genotypes (e.g. rCGS 41) are less
affected by the disease than is the commonly grown groundnut cultivar
55-437.
10.5.5 Seedborne nematode diseases

Aphelenchoides arachidis Bos and Ditylenchus destructor Thorne infect the
seeds of groundnut. Aphelenchoides arachidis distribution is limited to a
few isolated areas in northern Nigeria (Bos, 1977). Ditylenchus destructor
is reported from major groundnut-producing regions of the Republic of
South Africa and can destroy 40-60% seeds in heavily infested fields (De
Waele et al., 1988). These diseases have not been reported on groundnut in
other parts of the world.
A. arachidis is a facultative endoparasite of groundnut and it parasitizes
tissues of pods, testas, roots and hypocotyls, but not the cotyledons,
378 Diseases
embryos or other parts of the plant (Bos, 1977; Bridge et at., 1977). Testas
infested with A. arachidis are thicker and more uneven than normal.
Heavily infested seeds are light brown; examined at lifting, their testas are
translucent and have dark vascular strands within them. Testas of the
infected dried seeds are dull in colour and often are wrinkled. Nematode
damage has little effect on seed germination.
D. destructor is found in roots, pegs, shells and seeds (de Waele et at.,
1988). The initial symptom of the disease is development of brown necrotic
tissue at the pod base, at the juncture of peg and pod. Infected pods lack
the lustre of healthy pods. Infected seeds are usually shrunken and the
micropyles are dark brown to black. Infected embryos are usually olive
green to brown.
Very limited information is available on the extent of losses caused by
these nematodes, their interrelationships with other micro-organisms and
their control (Bridge et at., 1977; Minton and Baujard, 1990).
No groundnut cultivars resistant to either A. arachidis or D. destructor
have been reported. Immersing seeds infested with A. arachidis in four
times their volume of water and heating at 60°C for five minutes gives
complete control of the nematode without adversely affecting germination.
Sun-drying of pods after harvest in very dry conditions reduces the number
of viable nematodes in the pods (Bridge et at., 1977). One method of
managing D. destructor is rotation of groundnut with non-hosts and less
susceptible crops. Maize and grain sorghum crops should not precede
groundnut as a rotational crop in fields infested with the nematode. Wheat
is apparently a poor host (Basson et al.,1990). Unless appropriate pre-
cautions are taken, the seedborne nematodes could be disseminated with
seeds and become serious pests worldwide.
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Published as ICRISATS Publication 1.A. No 1468

CHAPTER 11
Groundnut pests
].A. Wightman and G. V. Ranga Rao
11.1 INTRODUCTION
There have been four major reviews of the literature discussing the insects
living on groundnut plants since 1973 (Feakin, 1973; Smith and Barfield,
1982; Wightman et al., 1990; Gahukar, 1992). There are also several more
concise accounts dealing with pest problems in general (e.g. Wightman and
Amin, 1988; Wightman et al., 1989; Lynch and Douce, 1992), specific
topics, such as host plant resistance (Lynch, 1990), and discrete geographi-
cal zones such as India (Amin, 1988), developed countries (Biddle et at.,
1992) and southern Africa (Wightman, 1988a; 1989; Sohati and
Sithanantham, 1990; Sithanantham et at., 1990). Feakin (1973), Redlinger
and Davis (1982), Dick (1987a,b) and Wightman et al. (1990) provide
details of the insect pest problems associated with stored groundnut and
their management in developed and developing countries. There is little
more to add to what has already been recorded about the post-harvest
pests of groundnut - the limited coverage given to them in this chapter
should not be taken as an indication that they lack importance. Thus,
although the general literature up to 20 years ago was somewhat sparse
(despite the publication of a large body of information in primary sources),
there have since been attempts to redress the situation.
Smith and Barfield (1982) extended Feakin's (1973) pioneer work by
providing an invaluable list of the pest species associated with the crop.
There is not yet enough verified or verifiable data to distinguish between
those insects that merely live on, under or around groundnut stands
without causing appreciable damage, and those that are capable of causing
significant or economic reductions in crop yields when their populations
reach a particular intensity. We call the latter pests, restricting the term to
mean 'insects' as opposed to all biotic constraints. For the sake of simplifi-
cation, we include all yield-reducing arthropods (in particular, myriapods
and arachnids) with the insects. The term intensity is adopted to indicate
that yield loss can be influenced by pest density, the duration of the
396 Groundnut pests
exposure of a crop to a cohort of insects and the combination of the two.
Insect-days is a convenient measure of insect intensity.
11.1.1 The taxonomic status of groundnut pests

Smith and Barfield's list includes more than 360 species of insect that were
known to be associated with the crop in the early 1980s. It could be
assumed that this list could be extended considerably after 20 years and
that it would be appropriate to include an updated version in this chapter.
However, this has not happened because progress is uneven. Lists that
supplement Smith and Barfield (1982) are included in this chapter where
specialized input has made them available.
The slow progress in extending our knowledge of the groundnut fauna is
partIy attributable to the fact that many groundnut-growing areas have not
been systematically surveyed for insects, especially in Asia and Africa.
Secondly, there remains the problem of locating taxonomists and paying
for the identification of the insects located during surveys (Wightman,
1988b). This is likely to be a major stumbling block because there is reason
to believe, as is explained below, that significant sectors of the all-
important soil fauna of the agroecological zones involved have not been
described. Thus even if surveys are carried out, many of the insects
collected may be new to science. The same problems - lack of knowledge
and lack of experts - are associated with the natural enemies of the
potential pests. Unfortunately, the trend for the withdrawal of the public
money needed to support prestigious centres of insect systematics does not
give hope that the situation will improve in the future.
It is admitted that it is not necessary to attach a latin binomial to an
insect to know that it is destroying part of a farmer's crop. Indeed the taxa
(above the level of genus) most likely to be damaging groundnut crops are
well known and are described briefly in section 11.2; Wightman et at.
(1990) give a more extensive account. However, a certain amount of
information is needed about distribution and the biology of insects before
management strategies can be developed. Pest species often coexist with
closely related, but harmless, members of the same taxon. Specialist input
from taxonomists is needed to ensure that non-taxonomists are able to
distinguish pests and non-pests at all stages of their life-cycles, preferably
in field conditions. These comments reflect a theme that will reappear
elsewhere in this chapter: what do we (still) need to know?
11.1.2 Where are the groundnut entomologists?

Three sectors have generated and collated the information about manage-
ment of groundnut pests: university and government scientists of the
national agricultural research systems (NARS) of the countries in which
groundnut is important, bilateral research agencies, and the Consultative
Group for International Agricultural Research (CGIAR).
Introduction 397
The lines of demarcation are blurred. For instance, entomologists from
the US Department of Agriculture and two state universities have been
part of a bilateral programme of the Peanut Cooperative Research Support
Program (P-CRSP) in south-east Asia and West Africa. This has involved
American scientists working in the target countries, as well as scientists
from less developed countries visiting the USA to tackle some of its
endemic peanut research problems. The International Crops Research
Institute for the Semi-Arid Tropics (ICRISAT), which is based in peninsu-
lar India, is the CGIAR Institute with a mandate including groundnut. The
current focus of its two groundnut entomologists is almost entirely on Asia
but the ICRISAT Center in India accommodates many scientists from
elsewhere, including entomologists from other centres of excellence, such
as the United Kingdom's Natural Resources Institute (NRI), to work on
mandate-related problems of mutual interest. Much of the germplasm
tested for pest resistance by the American scientists in Africa and Asia was
supplied through the Genetic Resources Unit at ICRISAT. However,
many of the test lines originated in US laboratories and came originally
from the farms of many countries. We freely and gratefully acknowledge
the international effort that lies behind the production of much of the
information contained in this chapter.
A considerable amount of new information has become available in the
last 10 years or so. Some of it is had not been published formally at the time
of writing (1993) and is therefore presented in more depth than would
normally be the case, with due reference and deference being made to its
originators.
11.1.3 Emphasis
There is a bias in this chapter towards emphasizing information that is
available or that is needed to develop pest management systems for the
ground nut farmers of less developed countries. Stress is given to infor-
mation that is needed to understand the ecology of the insects (section
11.3) and the application of the pest management tools that farmers have
at their disposal (the subsequent sections). The next step is to evaluate how
effective these tools are, their relationship with other aspects of farm
management and how they may be handled to support the farming com-
munities. This is discussed in the final section.
Groundnut insects are not unique just because they live on this rather
special crop. We have therefore drawn on examples related to other crops
and insects to illustrate specific points.
11.1.4 Integrated pest management: scientists and farmers

This chapter leans towards discussing groundnut pests from the point of
view of integrated pest management (IPM). This concept is seen as a
398 Groundnut pests
general move away from complete reliance on synthetic insecticides for
managing (i.e. killing) insects that may have a detrimental effect on crop
yields. There may also be a case for suggesting changes to the management
of the many farms that grow groundnut without insecticide application.
The alternatives are combinations of:
• the provision of crop varieties that are in some way resistant to the most
important pest(s);
• sowing crops in combinations or patterns that result in pest outbreaks
being diminished in intensity and frequency: such modifications may
involve the physical environment, and/or
• changes in the farm environment that encourage the natural enemies of
potential pests to aggregate within the cropping system;
• enhancing natural control processes by releasing biological agents such
as specific insect pathogens, parasites or predators in the farm
environment;
• applying natural or synthetic insecticides only when they are needed and
in such a way that the impact of predators and parasites is not
diminished.
A common feature of these intertwined alternatives is the emphasis on the
prevention of pest outbreaks. It is also clear that they are not necessarily
crop specific, and not always even farm specific. There can be good reasons
for IPM being the business of the community.
These concepts are the basis of the IPM tool box and involve making
management decisions beyond the traditional or conventional gamut of
what has been known as pest management. The people who are going to
make these decisions are the farmers. The role of the scientists (of several
disciplines) and extension workers is to evaluate procedures that may not
have occurred to farmers and to demonstrate alternative approaches to
procedures that the advisors see as being detrimental. Farmers in develop-
ing countries can rarely employ specialists (soil analysts, IPM scouts,
disease forecasters, etc.) to help them make decisions. Thus, although a
team of scientists can and should ensure that IPM procedures 'fit' particu-
lar sets of environments, it is the farmer who has to make the day-to-day
and season-by-season decisions according to his or her own perceptions of
the resource base of the farm or family, its requirements and aspirations.
Once they have their initial data sets, scientists genuinely interested in
IPM should probably consider the need for interaction between the labora-
tory and the land, the farmer, the advisor and the researcher. General
principles can be worked out on research stations but do not gain relevance
until they are established as being viable on a number of farmers' fields.
However, IPM is knowledge intensive, and some of the available knowl-
edge is presented below.
The insects 399
11.2 THE INSECTS
This section describes the insect taxa most likely to be associated with
reduced groundnut production, together with an indication of their distri-
bution and the kind of damage they cause. Further details of their biology
and ecology are available in Wightman et al. (1990) and in the other
publications indicated below.
11.2.1 Soil insects

The insects that live in the soil of groundnut fields are responsible for
higher levels of yield loss than foliage feeders. They attack pods and roots,
and the foliage via the roots. Anitha (1992) has constructed a key to the
pod borers based on damage symptoms.
Soil insects are difficult to manage because farmers usually do not know
that they are present until plants die or until the crop is harvested. The
prophylactic application of insecticides at sowing is not generally a feasible
proposition because of the non-availability of suitable products in many
countries, their high cost, and the residue problems many create in the
seed. The latter point refers specifically to the lipophilic but highly
effective cyclodienes such as dieldrin, aldrin, endrin and heptachlor. The
unacceptability of organochlorine residues in the environment in general,
and in the oil of groundnut seeds in particular, has meant that farmers have
lost access to the only persistent insecticides that give good control of soil
insects at a low price. However, they did present a risk to the health of the
applicators and of other non-target organisms. Progress is being made in
other areas of soil pest management following international recognition
that a widespread problem exists.
(a) Isoptera - termites

Termites are pests of groundnut throughout Africa, and in western and
southern Asia. Several of the most troublesome species are distributed
throughout this rather large region. Their attacks are usually associated
with periods of drought and therefore tend to be most serious at the end of
the growing season. They are less serious in Asia as a whole because they
prefer sandy or at least light, well-drained soil. Their life style is thus not
compatible with the lowland paddy systems that dominate much of the
agricultural landscape of this continent. However, as Wood and Cowie
(1988) list three of the termite genera mentioned below as being pests of
upland rice in West Africa, it is apparent that the cultivation of this crop is
not a complete barrier to termites. They are also rated as being a major
groundnut pest in Nepal (personal observation) and are recognized as pests
in southern Asia and Thailand (Wightman et al. ,1990; Logan et al., 1992).
400 Groundnut pests
In addition to being post-harvest pests (section 11.2.3), termites deplete
stand density and damage pods and seeds. Microtermes spp., especially
obesi and lepidus, penetrate the root system and quickly cause plant death,
certainly by their own feeding activity but also by exposing the plant tissue
to soil pathogens. Root penetration is usually as a result of their own
burrowing but it can occur via the lesions left by white grubs, wire worms,
etc. Nasutitermes apparently causes similar damage to Microtermes.
Ancistrotermes latinotus also penetrates the roots, but is more likely to
attack closer to the crown than Microtermes and then tunnel upwards
through the parenchyma tissue of the stems. In India, we have also found
Odontotermes obesi, wallonensis and brunneus penetrating the roots of
groundnut plants, causing damage similar to (but distinguishable from)
that of M. obesus.
Odontotermes spp, (especially obesus and brunneus) carry soil up the
stems and leaves and remove living tissue from within the shelter of this
soil sheeting, which presumably protects the workers from solar radiation
and natural enemies.
In Africa, Macrotermes spp.and Hodotermes mossambicus feed on stems
at the level of the crown. They can 'fell' whole plants but more often cut
out up to about half of the stems before moving on to another plant, which
is not necessarily next to the one they previously attacked. The economic
importance of the so-called harvester termites is often underestimated
because their activities are likely to go unnoticed. The fallen stems quickly
disappear as a result of the activities of other insects, especially soil
sheeting termites such as Odontotermes spp.
Microtermes spp. also attack pods. Sometimes they penetrate the pods
and remove the seeds; more often, they 'scarify' the shell by removing the
corky tissue from between the veins of the pod. This weakens the shell
considerably, resulting in exposure of the seed and its damage during
harvest and subsequent handling. More importantly, scarification also
increases the probability that the seed will be contaminated by aflatoxins -
carcinogens produced by common soil fungi of the genus Aspergillus,
especially A. flavus.
(b) Dermaptera - earwig'>

A little more is now known about earwigs than was indicated by Wightman
et al. (1990). On the vertsiols (black or clay cotton soils) of the ICRISAT
farm in peninsular India, Euborella annulipes, E. plebeja and Forcipula
quadrispinosa (Anitha, 1992) can reach popUlation densities that result in
damage to more than half of the harvestable pods in a stand. No satisfac-
tory insecticidal control method is documented.
Pitfall trapping in Alabama revealed that predacious earwigs, Labidura
riparia, were the dominant species living on the soil surface in the peanut
The insects 401
fields studied over a two-year period (Kharboutli and Mack, 1991). Thus
not all earwigs are potential pests.
(c) Hemiptera - Homoptera

Pseudococcidae - mealybugs Mealybugs, such as the cosmopolitan
Pseudococcus solani, are frequently found on the roots of groundnut
plants. The comparatively small size of colonized plants is an indication
that they are capable of reducing per-plant yield and that they may be a
pest but further surveys are needed to establish their importance. The
pineapple mealybug, Dysmicoccus brevipes (Cockerell), is occasionally
found in Australian groundnut crops (Brier and Rogers, pers. comm.).
Tettigometridae - hilda(bug) Hilda patruelis is one of the most serious

constraints to groundnut production in sub-Saharan Africa. This is not only
because of the across-farm plant mortality it causes but also because the
uncontrollable outbreaks can turn farmers away from this crop for many
years.
It lives on the roots of groundnut plants but is equally at home on the
stems of some of its other hosts, which include common herbaceous weeds
and woody ornamentals. The eggs are parasitized by Psyllechthrus oopha-
gus (Hymenoptera: Encyrtidae) but the rate of attack falls off markedly in
the rainy season, which is when the groundnut crops are in the ground.
There is thus a case for obtaining a better understanding of the ecology of
the parasite. Outbreaks of hilda are most likely to occur when crops suffer
from drought stress, especially when farmers sow after pre-season showers
that are followed by a dry spell. This species is remarkably mobile for an
insect that is adapted to life underground.
Hypochthonellidae Hypochthonella caeca was recognized as a sporadic

pest of groundnut in Zimbabwe by Rose (1962). It has since been found on
groundnut roots in Zambia by Wightman. In a letter published in the
International Arachis Newsletter (No.9, May 1991, p. 4) Dr M.R. Wilson,
International Institute of Entomology, London, indicated that this species
is no longer considered to be a member of the Tettigometridae but is the
sole member of the Hypochthonellidae.
(d) Hymenoptera
Formicidae - ants Most of the ants encountered in groundnut fields are
predators, especially of termites and caterpillars, although many of them
tend aphids and other homopterans in exchange for honeydew. On balance,
the inevitable presence of ants in groundnut fields (at least in Africa) is, by
virtue of their role as predators, a good reason for not promoting the use of
insecticides in these farming systems (Wightman and Wightman, 1988).
402 Groundnut pests
The only species of ant known to be groundnut pests are Dorylus
orientalis and D. labitus (doryline, blind or red ants). The ants hollow out
the pods as they approach maturity, entering by regular holes 2-3 mm in
diameter.
Both species have been known as pests of groundnut in India and
Malaysia (Dammerman, 1929) for many years. Only in Thailand is the
problem sufficiently serious for action to be taken to manage Dorylus spp.
(poisoned coconut meat baits). It is recently believed to have been found in
Philippine groundnut fields (ICRISAT, 1991a). There is thus a good case
for a concerted search of groundnut fields in the countries between
the Philippines and India to determine its range and perhaps the site char-
acteristics that govern its presence or absence in Asia. Wightman and
Wightman have found Dorylus sp. under groundnut in Malawi, Zambia
and Zimbabwe and have detected pod damage in Malawi.
(e) Coleoptera
Buprestidae - jewel beetles The jewel beetle Sphenoptera indica, which is
a root borer, has been known as a widespread resident of groundnut fields
in India for many years (Rai, 1979). Information about its potential pest
status has recently become available (Logan et al., in press). In a rain-fed
field on the ICRISAT Research Farm 20 days before the 1986 rainy-season
harvest, it was found that 23% of the groundnut plants had S. indica in
their roots as larvae, pupae or adults. As the three larval instars tunnel
through the parenchyma of the root, a high rate of mortality or severe
wilting among the attacked plants was anticipated. On one side of the field,
where 76% of the plants were attacked, 82% of the dead plants and 64% of
the living plants were host to this species. A survey of groundnut plants
growing in three irrigated fields on the ICRISAT farm in the 1986 rainy
season revealed infestation rates of 0.4%,9% and 14%. This indicates that
irrigation does not eliminate the risk of attack by this species. Attacks in
subsequent years have been sporadic, and spread across the 1300 ha farm;
they have been sufficiently heavy to permit us to screen for resistance to
this species among Arachis spp. A survey in Andhra Pradesh and
Karnataka (southern India) in the rainy season of 1992 revealed that up to
10% of the plants growing on red soils (alfisols) had been killed by this
species.
Scarabaeidae - white grubs (as larvae); cockchafers, May bugs or June

beetles (as adults) The general importance of white grubs as pests of
groundnut in India and in parts of southern Africa has been recognized
only recently (Wightman et al., 1990). Since then, further research and
discussion with experts from Asian countries has revealed that white grubs
are associated with yield loss from India to China and the Philippines. They
are also a problem in moister areas of West Africa, such as Senegal.
The insects 403
Appert (1956) noted that 'underground larvae' (i.e. white grubs, Table
11.1) can cause as much damage as millipedes,which are usually recognized
as a major constraint to groundnut production in West Africa.
It has therefore been possible to extend the list of white grub species
associated with groundnut (Table 11.1; cf. Table 5.10 in Wightman et al.,
1990) to include further information from Asia and southern Queensland,
Australia, where the identity of the main peanut pest species has been
resolved and a key provided (Rogers et al., 1992).
It appears that there are some 70 named species (including subspecies)
of white grub known to be associated with the groundnut crop. However,
the current maximum exceeds 100 because of the number of undescribed
species. This compares with c. 52 species of termites (Wightman et al.,
1990) and more than 60 lepidopteran species (Smith and Barfield, 1982).
White grubs feed mainly on the taproots and/or the peripheral rootlets.
Either way, the net effect is to restrict the growth of the plant. This is
particularly marked in sandy soils in drought prone areas such as occur in
groundnut fields of the Middle Veldt of Zimbabwe. Seedlings can be killed
outright if the phenologies of the crop and pest result in large larvae and
small plants occupying the field at the same time. In the Sudan, white grubs
have been associated with aflatoxin contamination of groundnut left in the
ground for 6 weeks after it should have been harvested (Ahmed et al.,
1989).
Heteronyx piceus is the species most likely to cause damage in
Queensland. It forms 90% of the scarab population under groundnut in the
main growing area. It is unusual in that it attacks the pods but not the
roots. Population densities of up to 30 larvae per metre of row nave been
reported (Brier and Rogers, unpublished).
Kalshoven (1981) indicated that there are many species of Anomala in
the Indonesian archipelago. Supriyatin (1991) reported that white grubs
attack the roots of groundnut plants in upland crops in that country. As
members of this genus are known to eat groundnut roots in other
countries, it is likely that there may be an undefined white grub problem in
East and/or West Java, the centres of groundnut production in Indonesia.
Cadapan and Escano (1991) indicated that Leucopholis irrorata has
recently been associated with the groundnut crop in the Philippines. This
ties in with the major pest status awarded to this species in maize-rice
systems (Litsinger et al., 1983).
Asurvey by Nath and Singh (1987) of cropped fields (mainly groundnut
and sugar cane) in a relatively small area of eastern Uttar Pradesh,
northern India, added 16 species to the list. They indicated that these
species were common to all crops but were most numerous on groundnut
and sugarcane. Several of these species were also found during a detailed
study of white grubs many miles away in semi-arid Rajasthan, particularly
around Jaipur (Yadav, 1981). However, Yadav did not associate them
specifically with the groundnut crop in his report, which concentrates on
404 Groundnut pests
TABLE 11.1 Scarabaeidae associated with the groundnut crop in the larval (white
grub) stage
Species Location References
Adoretus cribrosus Zimbabwe Smith and Barfield, 1982

A. decanus India Nath and Singh, 1987
A. laisopygos India Nath and Singh, 1987
A.limbatus India Nath and Singh, 1987
A. lImbrosus 'Africa' Smith and Barfield, 1982
A. verstltus India Nath and Singh, 1987
Adoretus spp. Malawi, Zambia Wightman
(up to 4 spp.) Zimbabwe
Anomala antiqua Burma Smith and Barfield, 1982
A. atrovirens Indonesia Smith and Barfield, 1982
A. bengalensis India Nath and Singh, 1987
A. corpulenta China (PR) Xu, B.C., 1982
A. dorsalis India Yadav, 1981
A. dorsalis India Nath and Singh, 1987
var. fraterna
A. plebeja Senegal Appert, 1956
Burkina Faso IRAT,1976
A. punjabensis India Yadav, 1981
A. ruficapilla India Nath and Singh, 1987
A. rufocuprea Korea (Rep) Cho et al., 1989
Anomala spp. Botswana, India, Wightman; Yadav, 1981;
(up to 11 spp. in Indonesia, Malawi, Kalshoven, 1981
southern Africa) Zambia, Zimbabwe
Anomala sp. Burma R. Milner, personal
communication, not A. antiqua
Apogonia cribricollis India Nath and Singh, 1987
A. ferruginea India Nath and Singh, 1987
A. roucca India Nath and Singh, 1987
A. uniformis India Nath and Singh, 1987
Autoserica atratula India Nath and Singh, 1987
A. insanabilis India Nath and Singh, 1987
A. nathani India Nath and Singh, 1987
Dyscinetus trachipygus India Nath and Singh, 1987
Crator cuniculatlls Burkina Faso IRAT,1971
Eulepida mashona 'Africa' Smith and Barfield, 1982
Heteroligus claudius Nigeria Smith and Barfield, 1982
Heteronyx brevicollis Australia Smith and Barfield, 1982
H. piceus Australia Rogers et al., 1992
H. rugosipennis Australia Rogers et al., 1992
H. sp. nr. Australia Rogers et al., 1992
rugosipennis Australia Rogers et al., 1992
H. diomphalia China (PR) Shang et al., 1981
Korea (Rep) Cho et al., 1989
H olotrichia consanguinea India Smith and Barfield, 1982
H. formosana China (PR) Lu et al., 1987
The insects 405
TABLE 11.1 Cont.
Species Location References
H. morosa Korea (Rep) Cho et al., 1989

H. oblita China (PR) Wang et al., 1986
H. parallela China (PR) Wang et al., 1986
H. sauteri China (PR) Huang and Lin, 1987
H. serrata India Smith and Barfield, 1982
Lachnosterna caudata1 Australia Smith and Barfield, 1982
L. fissa India Smith and Barfield, 1982
Lepidiota sp. Australia Gough and Brown, 1988
L. crenita Australia D.l. Rogers and H.B. Briers,
personal communication
Leucopholis irrorata Philippines Cadapan and Escano, 1991
Maladera orientalis China (PR) Wang et al., 1986
Korea (Rep) Cho et al., 1989
Maladera sp. Thailand Sathorn Sirisingh, personal
communication
Neodon pecuarius Australia Smith, 1946
(= Trissodon
puncticollis)
Oxycetonia versicolor India Smith and Barfield, 1982
Pentodon idiota USSR Smith and Barfield, 1982
Phyllophaga ephilida 'Americas' Smith and Barfield, 1982
P. armicans 'Americas' Smith and Barfield, 1982
Podalgus (Crator) 'Africa' Smith and Barfield, 1982
cuniculus
Popillia japonica China (PR), N. America Smith and Barfield, 1982
Pseudoheteronyx basicollis Australia P.G. Allsopp, in Rogers et al.,
1992
Rhopaea magicornis Australia Smith and Barfield, 1982
Schizonycha africana NE Africa Smith and Barfield, 1982
Senegal Appert, 1956
S. fusca Malawi Wightman
S. ruficollis India Nath and Singh, 1987
S. straminea Malawi Wightman
Schizonycha spp. (up to 8) Malawi Wightman
Sericesthis ina Australia Rogers et al., 1992
S. suturalis Australia Rogers et al., 1992
Schizonycha spp. (up to 3) Malawi Wightman
Strigoderma arboricola USA Smith and Barfield, 1982
Trochalus pilula Senegal Appert, 1956
Trochalus sp. Malawi Wightman
Xylotrupes gideon Burma Smith and Barfield, 1982
Tribe: Sericini 8 indet. Malawi, Zambia, Wightman
spp. Zimbabwe
l. not recognized in Australia, possibly Lepidiota caudata, see Gough and Brown (1988)
(P.G. Allsopp, personal communication)
406 Groundnut pests
the predominant Holotrichia consanguinea. The predominant species in
southern India is H. serrata (Veeresh, 1977).
Elateridae - Click beetle larvae or wireworms; Tenebrionidae - false wire-

worms Wireworms and false wireworms can be treated together because
they have a similar morphology and the symptoms of their pod boring
actIVIties are indistinguishable: both make symmetrical holes,
2-3 mm in diameter, in the maturing and mature pods. Elaterid larvae
have also been discovered eating the tissues of germinating seeds on the
ICRISA T research farm in peninsular India. This activity could lead to
diseased and malformed plants.
Appert (1956) indicated that wireworms ('taupins'), mentioning
Cardiophorus subspinosus and C. cognatus, can be predators but, together
with false wireworms, they can reduce the density of seedling stands by as
much as 10%.
The survey carried out by Wightman in southern Africa in 1987 revealed
a major problem with both of these taxa: the difficulty of identifying them
to species from the larval stages and sometimes the adults. Table 11.2
shows species in the Elateridae and Tenebrionidae added by Wightman
(with determinations by the British Museum of Natural History) to those
listed by Smith and Barfield (1982).
Chrysomelidae Members of this large family will almost certainly be

found tasting the foliage of groundnut wherever it is grown, but they
probably cause little damage. The same cannot be said for members of the
genus Diabrotica, especially D. undecimpunctata howardi, the spotted
cucumber beetle, in the southern states of the USA. The adults cause
relatively minor damage to the terminal leaves but the subterranean larvae
can cause major injury to the pods and pegs. As such, it has been recog-
nized as a major pest of this crop throughout this century (Smith and
Barfield, 1982).
Curculionidae - Weevils The only species of weevil that has had a high
profile as a pest of groundnut is the white fringed weevil, Graphognathus
leucoloma (Feakin, 1973), especially in the Americas and, more recently,
in Australia. The larvae eat the roots and cause stunting of the stems and
plant death. The adults (parthenogenetic females) eat the foliage. Each
can lay 1000-2000 eggs, hence the high damage potential of this species.
Adult weevils of other species are often numerous in groundnut crops
and can often be seen eating the edges of the leaflets. In southern Asia, the
ash grey or grey cotton weevil Myllocerus undecimpustulatus maculosus is
often found at densities of up to 10 per plant. Systates spp., Mesoleurus
dentipes and Diaecoderus sp. can reach even higher densities in southern
Africa (Jepson, 1948; Rose, 1962; Broad, 1966). Jepson (1948) associated
S. articollis with the 'yellowing and failure' of a young plantation. This is a
The insects 407
TABLE 11.2 Additions to species lists (Smith and Barfield, 1982) in Elataridae
and Tenebrionidae
Species Locations
Elataridae (two previously listed)

Agrypninae, Agrypnini (larvae) indet. Mtopwa, Tanzania
Agrypninae, Monocrepidiini (larvae) indet. Mawengo, Zimbabwe
Elaterina (larvae) possibly 6 spp. indet. Malawi and Zambia
Tenebrionidae (five previously listed)
Pimeliinae (larvae) possibly 13 spp. From 20 locations in Malawi,
Zambia and Zimbabwe
Tenebrioninae (larvae) possibly 3 spp. Malawi and Zimbabwe
Zophosis sp. (adults) Malawi
Gonocephalum nr simplex (adults) Malawi
Anchophthalamus plicipennis (adults) Malawi
Drosochrus sp. (adults) Malawi
significant observation because it is likely that the larval stages of weevils

caused the yellowing (by attacking the nodules, cf. Sitona spp.) and crop
failure by eating the lateral roots. The importance of weevil larvae as
subterranean groundnut pests is worthy of further consideration.
(f) Lepidoptera
The caterpillars of several lepidopteran species that live at the soil surface
damage groundnut plants - Agrotis spp. and Feltia spp. feed at the crown,
Spodoptera litura (in India) and S. littoralis (in southern Africa) are pod
borers. In Australia, the larvae of Etiella behrii, the lucerne seed web
moth, penetrate the pods and feed on the seed (Brier, personal
communication) .
The only true soil-dwelling lepidopteran to cause major damage to
groundnut is Elasmopalpus lignosellus, the lesser corn stalk borer, which is
restricted to the New World (Smith and Barfield, 1982). This species is
regarded as a major pest of groundnut and other crops. Larvae feed at or
close to the soil surface in the first two instars, paying particular attention
to the flower and vegetative buds. The older stages feed on the under-
ground parts of the plant, and often scarify the pods. This results in a high
risk of seed contamination with aflatoxin (Lynch and Wilson, 1991).
(g) Myriapoda - millipedes

Millipedes, often more than 30 cm long, are the most important pests of
groundnut in the drier areas of West Africa, where they attack the pods,
408 Groundnut pests
mainly before they have hardened (ICRISAT, 1988). They are also recog-
nized as pod borers of lesser importance in southern Africa, where they
damage 5-10% of the immature pods, some of which would not reach
maturity by harvest time (Wightman, 1989).
Their importance in West Africa is related to the amount of damage they
do, the unpredictable nature of the attack and because there is no control
method available, irrespective of the socio-economic status of the farmers
who have to contend with them. The literature reviewed by Wightman et
al. (1991) indicates that the impetus of research carried out on the ecology
and control of millipedes in the 1970s has not been maintained.
11.2.2 Insects that live on the leaves and flowers
(a) Orthopteroid orders - grasshoppers, locusts, crickets, mantids

Orthopteroid insects are frequently found in groundnut crops but, with the
exception of locust plagues, no record of them achieving pest status has
been located. Observation indicates that their presence in groundnut
foliage is associated more with sunbathing than feeding. Mantids appear to
be an exception. They can achieve relatively high densities in southern
Africa (Wightman).
(b) Thysanoptera - thrips

Smith and Barfield (1982) list 18 species that have been associated with
groundnut crops. Several species can be added to this list:
Megalurothrips usitatus is frequently encountered in the flowers of ground-
nut in Asia and southern Africa (Palmer et al., 1990; Wightman et al.,
1990).
Scirtothrips aurantii was detected in terminal (folded) leaflets in Malawi
(Wightman). S. oligochaetus has been found on groundnut in India
(Palmer et al., 1990).
Thrips palmi has been found in the terminal leaflets of groundnut in India
and other Asian countries (Palmer et al., 1990) and also lives in Australia
and the 'Pacific' but has not been recorded from New Zealand and New
Guinea (Houston et at., 1991). T. setosus is known only from Japan and
Korea (Reddy et al., 1991).
Each species has a preferred niche - presumably the feeding site - within a
plant. However, flower dwellers can be found in other parts of the plant
before flowering occurs (Table 11.3).
Thrips can have pest status in groundnut crops as virus vectors and as
leaf-eaters. We believe that thrips are also of considerable pest status
because the (largely cosmetic) damage they cause induces farmers to apply
insecticides unnecessarily (Lynch et al., 1984; Ranga Rao and Shanower,
The insects 409
TABLE 11.3 The distribution of thrips between flowers and leaf buds in the rainy
and post-rainy seasons at fCRfSA T Center, 1990-1991 (pooled data)
Total number of thrips observed (% in parentheses)
Rainy season Post-rainy season
Leaf Flower Leaf Flower
Scirtothrips dorsalis 9366 455 5274 153

(97) (26) (85) (6)
Thrips palmi 148 102 792 361
(1.5) (6) (13) (13)
Frankliniella schultzei 145 1202 118 2183
(1.5) (68) (2) (81)
1988; Reddy et al., 1991). This results in outbreaks of other pests because
of interference in the natural insect density control processes.
Thrips as virus vectors

Thrips can have pest status in groundnut crops as virus vectors and as leaf-
eaters. They are vectors of the tomato spotted wilt virus (TSWV) and its
Indian variant, the bud necrosis virus (BNV). The various isolates of this
disease can cause widespread damage to ground nut in many countries
(Reddy et al., 1991).
Not all groundnut thrips are virus vectors. In India the most important
vector is T. palmi (not Frankliniella schultzei, as was previously reported,
although this species can be forced to transmit at a very low rate in
laboratory conditions). Scirtothrips spp. are not vectors in India (Palmer et
al., 1990; Vijayalaksmi pers. comm.).
The most likely vectors in the southern USA are F. occidentalis and
F. fusca (Reddy et al., 1991; Culbreath et al., 1992). Although T. tabaci is a
vector of TSWV in other crops, it has not been linked with outbreaks of
this disease in groundnut crops in the USA or elsewhere (Reddy and
Wightman, 1988; Reddy et al., 1991).
Thrips as leaf-eaters
Thrips can cause damage as a result of their feeding activity. The leaf-
eating species are usually found between the folded leaflets at the stem
tips. They cause little visible damage at the time of feeding. As the leaflets
grow, the small lesions and patches of dead cells left by the thrips do not
expand at the same rate (if at all) as the undamaged cells. This means that
the most conspicuous leaflets at the top of the plant are contorted and have
small holes in them. This damage is most apparent in young plants when
410 Groundnut pests
the temperature is not high enough to promote rapid growth, e.g. during
spring in North America and the post-rainy season in India. The most
acute case is that of Enneothrips flavens in Brazil. Increases in yield
following insecticide application and attributed to the control of this
species range from 35% to 50% (Smith and Barfield, 1982).
A debate about the economic status of thrips (mainly T. fusca) in
southern USA seems to have reverberated around the peanut industry for
many years (Lynch et al., 1984). Turnjit (1988) added further light to the
situation by indicating that in North Carolina no yield loss will occur until
40-50% of the leaflets are damaged, according to the variety, but then only
in plants less than 4 weeks old.
The situation in India is not clear, mainly because the populations of the
main leaf damagers (F. schultzei and S. dorsalis) cohabit with jassids
(Empoasca kerri) and there is no method of separating out the relation-
ships between density and yield losses attributable to each taxon. The
impression is that thrips cause little yield loss, at least on the ICRISAT
farm.
(c) Homoptera
Aphididae - aphids The aphid species most frequently linked with
groundnut is Aphis craccivora, the groundnut or cowpea aphid. It occurs
throughout the tropics and subtropics and has many hosts. It normally
appears on groundnut crops in the early rainy season, when it can cause
considerable damage to young plants. In recent years a second outbreak
has occurred in the post-rainy season on the ICRISAT farm and this may
be linked with a perceived air pollution problem (Dohman et al., 1984).
This species is of particular significance because it is the vector of the
(persistent) groundnut rosette virus complex in Africa. This disease can be
crippling but has become less common in recent years, especially in
southern Africa, where the widespread adoption of regulatory cultural
practices (especially earlier and denser sowing than was once practised)
may have limited its potential effects.
A. craccivora is capable of reducing the yield of groundnut crops by
means of its feeding activity alone (Mayeux, 1984; Bakhetia and Sidhu,
1976). Our experience in southern India indicates that populations are
regulated by coccinellids and other predators and rarely survive a spell of
persistent rain.
As representatives of the aerial plankton that overflies agricultural
areas, many 'other species' - such as A. robiniae, A. gylycines, A gossypii,
A. solanella, Myzus persicae, Macrosiphum euphorbiae, M. avenae,
Rhophalosiphum padi and Lipaphis erysimi - are implicated in the non-
persistent transmission of most other groundnut virus diseases, including
peanut stripe, cucumber mosaic, mottle and peanut stunt viruses. A.
gossypii transfers groundnut streak necrosis disease (= sunflower yellow
The insects 411
blotch virus) to groundnut in the Rift Valley area of southern Africa from
Tridax procumbens, a common tropical weed (Saleh, 1991; Wightman et
al., 1990; Xu Zeyong etal., 1991).
Cicadellidae ar lassidae - jassids ar leaf-happers Jassids are a common

feature of groundnut crops in most parts of the world. Perhaps they are so
familiar that entomologists have not bothered to collect them, because
only 20 species have been associated with the groundnut crop (Smith and
Barfield, 1982; Wightman et al., 1990). The economically significant genus
Empaasca predominates - E. fabae in the Americas, E. kerri in India - but
also lacabiasca farmasana in south-east Asia (but the latter two inadver-
tently omitted or not clearly referred to by the aforementioned authors)
and Austroasca alfalfae in Queensland. The symptoms commonly include
pronounced chlorosis followed by peripheral necrosis, a condition known
as hopper burn.
There are no definitive accounts of the relationships between their
density and crop yield. The main reason for this is the almost inevitable
cohabitation with thrips and other insects in the early stages of crop
development. The consensus in the USA, where experimentation has been
in progress for about 50 years, is that the damage caused by these insects is
cosmetic. Smith et al. (1985) indicated that the range of reduction in
photosynthetic area among 14 groundnut genotypes was 3.8-28%. This is
unlikely to have a direct effect on yield in view of groundnut's high leaf
area index (> 5 after 40 days). However, the shading of the lower leaflets
by damaged leaflets, which are usually on top of the canopy, may have a
greater effect than anticipated.
Aleyrodidae - whiteflies Feakin (1973) does not mention whiteflies and

Smith and Barfield (1982) indicate, correctly, that Bemisia tabaci is cosmo-
politan but add no more. This confirms our suspicion that the observed
outbreaks of whiteflies on groundnut in India and southern and south-
eastern USA are a recent phenomenon (Shanower and Ranga Rao, 1988,
Lynch and Simmons, 1993).
Bemisia tabaci epidemics have created problems associated with the
overuse of insecticides in cotton crops in coastal Andhra Pradesh. The
appearance of this insect on groundnut is considered to be associated with
this general problem but may have been created by the local overuse of
insecticides in groundnut. The situation is being monitored in view of the
ability of this species to debilitate crops and spread virus diseases.
The situation appears to be rather different in the USA where the cotton
strain (biotype A) has been replaced by the poinsettia strain (biotype B),
which is capable of defoliating groundnut plants. This was first recognized
in 1987 in Florida. Since then, the pest status of this species on groundnut
has worsened as populations of the 'new strain' have increased in density
and spread through Georgia and Texas (Lynch and Simmons, 1993).
412 Groundnut pests
TABLE 11.4 M iridae associated with the groundnut crop in addition to those listed
by Smith and Barfield, 1982
Species Location Source
Creontiades pallidus Lilongwe, Malawi J.A. Wightman (collected by

sweep net, 1987;
det. M.R. Wilson, eIE)
Creontiades sp. Queensland, Rogers and Brier (personal
Australia communication)
Creontiades pallidifer New Delhi, India Singh et al., 1990
Helopeltis sp. Lilongwe, Malawi J.A. Wightman
?Taylorilygus sp. Lilongwe, Malawi J.A. Wightman
(d) Heteroptera
Miridae Mirids can often be found in ground nut crops at low densities -
perhaps less than one per plant. Smith and Barfield (1982) list 11 species
belonging to eight genera from Africa, India and the USA. Table 11.4
shows additions to that list.
It is not often realized that this taxon can cause considerable damage at
low densities. For instance, in an Australian glasshouse experiment set up
in Queensland to demonstrate this point, Rogers and Brier (personal
communication) found that two adult Creontiades sp. per plant reduced
flower production by 86% over a 3-week period. Peg initiation showed a
corresponding decrease of 87%. The plants resumed flower production 7
days after the mirids were removed, and produced more flowers and pegs
than the unexposed control plants during the subsequent 4 weeks.
However, it is envisaged that compensation is unlikely to be possible under
the conditions of sustained attack that are likely to be encountered in field
conditions, so that there are serious implications in finding mirids in a
groundnut field. This is especially so in dryland agriculture where the
cohort of flowers that produces the main crop results from a particular
rainfall event that may not be repeated.
Damage in the Queensland experiment was typical of mirid attack to
legume crops in that the flower buds were attacked at their earliest
appearance and quickly became necrotic (Sorenson, 1936; Wightman and
Whitford, 1982; Clifford et al., 1983). The vegetative buds and other tissue
were apparently not attacked in this way so that there was no distortion of
the stems and leaves.
The potential of this species to cause damage is demonstrated by Singh et
al. (1990). The mirid populations that they detected on groundnut in New
Delhi peaked at just over one per plant in the rainy seasons of 1987 and
1988 and more than two per plant in the intervening summer crop. This
species was present from the second week after sowing until about the pod
maturation stage. It is unwise to transfer the conclusions derived from the
The insects 413
data of Rogers and Brier working in a glasshouse in Australia to field

conditions in India but we suggest that the mirids detected in New Delhi had
a marked effect on the rate of flowering and the subsequent yield. Clearly
the mirids living in groundnut crops need to be looked at rather closely.
Pentatomidae and Lygaeidae Members of these families are often con-

spicuous in groundnut crops but we have not attributed yield loss to the
'big bugs'. Their feeding activity appears to be concentrated on the vegeta-
tive tissue, especially the growing points, which take on a limp appearance
- hence the common name 'tip wilters'. Groundnut entomologists can
ignore them, unless they are found to be attacking the reproductive tissues.
(e) Lepidoptera
There are many species of leaf-eating caterpillars found on groundnut
plants - Smith and Barfield (1982) list more than 60 - and no doubt a
concerted search would find many more. Of these, relatively few are of
economic importance or limited to the groundnut crop. We wish to play
down the importance of defoliators to groundnut crop production because
natural control processes usually keep them at densities well below the
economic threshold. However, apparently spontaneous flare-ups can occur
and these are usually associated with the injudicious use of insecticides or
other examples of suboptimal management.
Spodoptera spp. (the armyworms) are prominent in the list of potential
pests, with S. frugiperda, ornithogalli, latifascia, sunia and eridania predo-
minating in the New World. S. littoralis and exempta are associated with
the groundnut crop in Africa and northern Asia, while S. fitura extends
across the remainder of Asia. S. exigua is cosmopolitan.
The heliothine genera can also be pests of groundnut over most of the
crop's range, e.g. Helicoverpa zea in North America and Heficoverpa
armigera in Asia, Africa and Australia. H. punctigera also feeds on
groundnut in Australia but, unlike H. armigera (a flower and peg feeder),
it is primarily a defoliator (H. Brier, personal communication). The arctiid
hairy caterpillars Amsacta spp. and Diacrisia obliqua are sporadic defolia-
tors in southern Asia. They can appear in devastatingly high numbers in
newly emerged rainy season crops. They are polyphagous and, fortui-
tously, usually have one generation per year.
Gelechiids predominate among the leaf miners, rollers, webbers and
tiers. Aproaerema modicella (= A. nertaria, Stomopteryx subsecivella,
S. nertaria, Anacampsis nertaria and Biloba subsecivella), the groundnut
leaf miner (Mohammad, 1981; Shanower et al., 1993; Wightman et al.,
1990), causes widespread damage and is fairly cosmopolitan within Asia. It
is known as a sporadic but potentially devastating pest with up to four
generations in one crop cycle. It is oligophagous, with a clear preference
for leguminous species, especially soybean and groundnut. Other hosts of
414 Groundnut pests
commercial importance include lucerne, pigeon-pea, mung bean and lab-
lab. On groundnut, the first three larval instars are leaf miners: when it gets
too large for this habit it becomes a leaf tier (or leaf folder). It pupates
between the folded leaflets.
Anarsia ephippias (groundnut leaf webber) can be a pest in northern
India. It webs the growing points and its feeding activity on the younger
leaflets results in 'shot-holing' as the leaves mature (Bakhetia, 1977).
Acarina - mites
Smith and Barfield (1982) have 17 entries under Acarina, 10 of which are
for Tetranychus spp. They are potentially a world-wide problem. The high
reproductive rate and short generation time of these mites mean that they
have a high potential for rapid population increase if the natural control
processes are disrupted or if specific environmental conditions are severe.
This happens when fungicides reduce the effectiveness of an entomopha-
gous fungus, when inappropriate irrigation methods are applied and/or
when insecticides kill other natural enemies (Campbell, 1978; G.V. Ranga
Rao et al., 1990). In Australia, the peanut mite Paraplanobia sp. appears in
groundnut fields during periods of prolonged drought. Population densities
crash after heavy rainfall (H. Brier, personal communication).
11.2.3 Post-harvest pests

The ecology and management of the post-harvest pests of groundnut have
been discussed in full by Dick (1987b) and in Wightman et al. (1990). In
general, the storage pests per se and the approaches to their management
are common to many other products and so this aspect is not dwelt upon
here. The emphasis is on several pests of the post-harvest situation in the
groundnut crop that are particularly relevant to Africa and Asia.
Attention is drawn to the period after the crop is harvested and before it
is 'plucked' and stored or bagged pending transfer to the market. During
this time it is usually stacked or windrowed to allow it to dry in the sun and
wind. This stage is critical in the production of a crop with high yield and
good seed quality and one that will not become the origin of contamination
with aflatoxin.
Unfortunately, the groundnut crop is not exempt from insect damage
during this time. Termites can come from below and remove significant
proportions (perhaps 30-40%) of the seeds and as much hay.
Odontotermes spp. are the most conspicuous in this regard, especially in
Africa (Logan et al., 1990; Logan et al., 1992). Pod damage at this stage
can add to the risk that stored material will become infected with
Aspergillus fiavus, the fungal source of aflatoxin.
The other pest specific to the drying stage is the 'wang', Elasmolomus
sordidus (= Aphanus sordidus), a lygaeid, which can be found in surpris-
Applied ecology and economics 415
ingly high numbers within stacks of drying groundnut plants. All stages
feed on the drying seeds by penetrating the pods with their mouthparts.
The net effect is a marked decline in quality caused by the build-up of
moulds and a rancid taste associated with the oxidation of oils to fatty
acids. This insect can be transferred into storage structures, where it
continues to cause seed deterioration.
The third species of note is the groundnut bruchid Caryedon serratus,
which is restricted to groundnut as a post-harvest pest. The only other
known host of commercial importance is the tamarind tree (Tamarindus
indica), the pods of which are traded mainly within southern Asian com-
munities. Groundnut stored in unbroken pods is usually safe from attack
by most insects and diseases, except with this species. The eggs are laid on
the pod, through which the neonate larvae dig to reach the seeds.
Warehouse contaminations probably originate from field infestations.
Although this pest was dealt with routinely in the heyday of West African
groundnut exports, its significance in India has only come to light in recent
years (Dick, 1987a).
11.3 APPLIED ECOLOGY AND ECONOMICS
Section 11.2 indicates that there are many different kinds of insect living on
or under groundnut crops and that some are undoubtedly influencing the
yield of the crop. The taxa and sometimes the species most likely to be
reducing yield have been mentioned with more or less detail. The sectors
of the many communities who have a stake in these matters (rich farmers,
poor farmers, female farmers and mothers, teachers, agroindustrialists,
extension specialists, researchers) will have a range of attitudes to these
insects according to their vested interests.
Some would take every opportunity to get rid of them with pesticides,
irrespective of their potential pest status; others would do so if they had the
means to purchase pesticides. Perhaps a few would ponder upon methods
of managing them without this drastic approach or would just like to have
the time to sit in a field and watch them. We are mainly concerned with the
section of this spectrum of interest that excludes the first category.
When trying to determine the status of the insects that live on a crop and
deciding whether they are pests, neutrals or beneficials, for instance, it is
necessary to have knowledge of their ecology and their influence on the
yield of the crop. This section reviews what is known about those aspects of
the applied ecology of the key pests that are fundamental to developing
what we regard as rational management strategies.
11.3.1 Flight activity

Most of the insects colonizing a crop originate from parental stock that
reproduced after flying into the field or its bordering vegetation. The same
416 Groundnut pests
is often true for virus vectors. Thus, when considering the risk of pest
attack, it is necessary to know about the times of year when economically
important insects are most likely to be flying, their pattern of movement
within and between localities, the relationship between flight intensity and
insect population density and yield loss, and the most effective methods of
monitoring their flight. This has a direct bearing on pest avoidance (section
11.7).
(a) Monitoring flight activity

The most effective method of catching flying insects is to lure a fraction of
the aerial population into a trap. Southwood (1978) discussed many
ways of doing this. Non-specific methods such as light traps (groundnut
leaf miner, Shanower, 1993), suction traps (for thrips, C.S. Gold,
ICRISAT, personal communication) and yellow water (pan-) traps for
aphids (Farrell, 1976b) have yielded reproducible results for insects living
in groundnut fields.
Even better are traps that are attractive to single species. With regard to
groundnut, this category is currently restricted to Lepidoptera, in particu-
lar Spodoptera litura (Ranga Rao et al., 1991a,b), S. frugiperda, S. exigua,
Helicoverpa zea, Heliothis virescens (Lynch and Douce, 1992) and the
groundnut leaf miner (ICRISAT 1991b) for which synthetic pheromones
are available. Traps enclosing virgin female moths have also been used to
lure male groundnut leaf miner moths but are considerably less effective
than the traps baited with the synthetic pheromone developed by NRI
(Chatham, UK) (Table 11.5; and Nandagopal and Reddy, 1990). Such
traps are satisfactory if it can be proved, or if it is accepted, that the catch
of males represents the activity of the population as a whole.
(b) Flight activity and its implications

The main flight period of Aphis craccivora, the vector of the groundnut
rosette virus, can be expected some 6 weeks after the first 'planting' rains in
southern Africa. This was the basis of the recommendation that groundnut
crops should be sown 'early'. This advice is tempered by the observation
that a crop, once it has been established for 40 days or more, is much less
likely to be adversely affected by this virus than are younger crops.
Adherence to the appropriate management practices is probably the
reason for the virtual non-appearance of this disease in southern Africa in
the last 20 years.
Ranga Rao et al. (1991a) compared the data from light and pheromone
traps set on the ICRISAT farm in their study of the flight activity of
S. litura. The former was the standard monitoring procedure until the mid
TABLE 11.5 Mean number of groundnut leaf miner caught in delta traps baited with experimental pheromone lures at ICRISAT Center
during rainy season, 1991
Mean number of moths trapped in different weeks of exposure
Lure type 1 2 3 4 5 6 7 8 9
'Small septa'
Fresh 696 545 650 245 50 69 74 34 23
Continuously exposed 654 703 630 360 58 70 78 22 14
'Big septa'
Fresh 922 800 856 468 72 108 137 38 22
Continuously exposed 990 804 573 326 32 30 22 34 19
Virgin female 195 185 40 84 11 52 14 9 6

Empty trap 7 20 7 5 2 16 3 1 3
SE ± 32.1 ± 37.8 ± 64.6 ± 41 ± 8.1 ± 13.9 ± 15.7 ± 7.3 ±3.4

CV(%) 12 17 31 37 48 62 64 71 51
418 Groundnut pests
TABLE 11.6 Relationship between log of numher of
moths caught (x) and log of number of egg masses (y)
per 100 plants (Kumari, 1989)
Site y [2
0.019 + 0.445x 0.798

2 -0.220 + 0.535x 0.826
3 -0.070 + 0.474x 0.838
x = mean of three traps per site and two nights' data combined
Data cover 38 observation periods between I February and 16
April 1989.
1980s. The pheromone traps consistently demonstrate a distinct peak of

flight activity in March (just before harvest of the post-rainy season crop)
that was not present in the light trap data. However, the implications are
not clear because there was no corresponding increase in oviposition in
groundnut crops at that time. This suggests that the females accompanying
the males were 'flying through' the farm. Perhaps they did not recognize
groundnut as a suitable host, or they may have been undergoing a long-
distance dispersal flight and were not attuned to respond to a potential
food plant.
In addition, the pheromone trap catches indicated that there was no
clear cessation in flight activity during the hottest time of the year. This
suggests that a period of aestivation cannot be assumed to be a normal
event, as suggested by the light trap catches. A periodicity of flight activity
corresponding to the generation length was also detected. This could be of
relevance to the timing of sowing in S. litura endemic areas, as it is best not
to sow at a time that will result in the colonization of the seedlings, which
tend to be the only stage susceptible to this pest.
One such endemic area is the coastal strip of Andhra Pradesh, India,
where groundnut is grown under irrigation from November to March.
S. litura is the only insect pest during this period. It can cause extensive
defoliation but this appears to be largely as a result of the disruption of
natural control processes associated with excessive insecticide application.
This area has been targeted as a test area for implementing IPM pro-
cedures based on damage forecasting via a pheromone trap network (Table
11.6) .
Kumari (1989) also demonstrated similar relationships between moth
catch and the density of small larvae (instars 1-3), large larvae and the
number of damaged leaflets per 100 plants. There was a delay of 4 days
between the appearance of the first moths in the traps and the detection of
the first egg masses. The delay was 8 days for the small larvae and 20 days
for the large larvae.
It is hoped that similar relationships will be established for the ground-
nut leaf miner now that a synthetic pheromone is available and trap
technology has been made effective (Hall, Cook, Ranga Rao and
Wightman, unpublished). Attention will be directed towards the establish-
ment of determining relationships between the number of male moths
caught per trap and the density of larvae in the next generation. The
feasibility of mating disruption exercises can then be investigated.
The groundnut leaf miner has not been observed to undergo long
migratory flights. Our observations on the ICRISAT farm indicate that it
will move only a matter of 50 m from a high concentration area (for
instance, in a soybean crop) to colonize groundnut. This means that
distributions can be extremely uneven, even'within a field.
Cockchafers (May beetles) are the adult stage of white grubs and have a
species characteristic flight pattern that is related to mating, feeding and
dispersal. The typical pattern is for adults to emerge at dusk over a period
of about 3 weeks or more at a precise time related to the time that the sun
goes over the horizon. First emergences occur over a period of 5-10 days.
Individuals do not appear every night; the females presumably spend a day
or so laying eggs in chambers, 10 cm or more below the soil surface (Farrell
and Wightman, 1972). The median first emergence date may be related to
temperature per se (day degree accumulation), a period of chilling
followed by warming in temperate climes (Wightman, 1974) or, in the
tropics, the onset of the rainy season (Yadav, 1981, 1991). In varying
combinations and sequences, according to the species, weather conditions
or the locality, the beetles:
• emerge from the ground (usually males before females);
• mate on the soil surface or on low-lying vegetation;
• undertake significant dispersal ('beeline') flights in a straight line for
>50 m away from the original emergence hole;
• feed on surface vegetation;
• fly towards and aggregate around markers, such as tall trees or telegraph
poles, and seek mates;
• fly to trees of a small range of species, and feed and mate on their
foliage;
• either drop off or flyaway from the tree when satiated or cold and, in
the case of females, lay eggs wherever they land.
This stage in the life-cycle of the Scarabaeidae is of economic importance
because it is during this period that the population is at its lowest density
and greatest accessibility to humans, i.e. it is the best time to attempt a
control strategy, if one can be devised.
Data sets on the host preferences and behaviour patterns of the adults
are needed. This information would, for instance, guide farmers as to
which tree species should be avoided or selected (as attractants or repel-
lents) in farm-forestry projects in areas where white grubs pose a high risk.
420 Groundnut pests
TABLE 11.7 Degree-days (0 D) required for completion of development from eggs
through to pre-oviposition for s. litura (Ranga Rao et ai., 1989) and groundnut leaf
miner (Shanower et ai., 1992a)
Stage S. litura a Groundnut leaf miner
°D Lower threshold
temperature (0C)
Eggs 64 60.1 (12.3)

Larvae 303 327 (8.9)
Pupae 155 72.3 (14.7)
Pre-oviposition period 29
Total
Range 508-631
Mean (543)
a Bred from egg masses found on the ICRISAT farm.

Temperature summation estimated from laboratory experiments carried out under constant
temperatures.
11.3.2 Environmental factors

Environmental factors can work directly on the insect:
• Temperature regulates the development rate and can be a cause of
mortality.
• Rain washes aphids off plants and promotes the high humidity that
stimulates entomophagous fungi.
• Wind influences the stimulus, distance and direction of migratory
flights.
The physical environment can have many indirect and often complex
interactions via the soil and, through the soil and its water or nutrient
content, to the plant and then the insect. Similarly, interactions between
temperature, relative humidity and cloud cover or levels of ultraviolet
radiation have observable effects (but without parameters) on the viru-
lence of insect pathogens.
(a) Direct effects

One of the basic sets of information that should be available for key pests is
the relationship between its development rate and temperature (Table
11.7). This is needed for matching insect damage or population models
with plant or crop development models. Biotype differences can also be
detected by comparing the relationship between temperature and the
development rate of isolated or transient populations.
Light, well-drained soils, as opposed to heavy, waterlogged soils, favour
the activities of Elasmopalpus lignosellus, especially when the weather is
hot and dry. Similarly, white grubs in general prefer sandy soil (Smith and
Barfield, 1982). An exception is Holotrichia serrata, which is most likely to
be found in the heavier soils of southern India, in contrast to its northern
counterpart, H. consanguinea, a denizen of the light soils of the Gangetic
Plain.
The opposite pertains for the southern corn rootworm (Diabrotica unde-
cimpunctata howardi). It proved particularly susceptible to dry soil, espe-
cially in the egg and first instar stage and when living in a sand medium
(Brust and House, 1990).
(b) Indirect effects

Wheatley et al. (1989) investigated the response of groundnut to drought
stressed hosts during the post-rainy season at ICRISAT Center. Four
varieties of groundnut were grown across a drought stress gradient. The
groundnut leafminer was most abundant on the most stressed plants.
lassids showed the reverse trend and this preference for a non-stressed
host was also observed for jassids living on lucerne (Hoffman and Hogg,
1991). Thrips were at first most abundant on the least stressed plants,
reversed this trend in mid-season and then reversed it back again before
harvest, by which time the physical condition of the hosts had deteriorated.
A further study might well investigate more closely the relative abundance
and feeding site of the thrips species involved. Of the four varieties tested,
NC Ac 343 (ICG 2271) proved to be the best to grow in times of potential
drought stress vis-ii-vis insect attack.
Several other aspects of this study were also of note:
• It was found that the groundnut leaf miner was able to withstand a
midday canopy temperature of 47°C.
• The source of the irrigation water - overhead or furrow - did not
influence the distribution of the insects.
• An inspection of the bamboo plot pegs indicated that termites (Micro-
termes sp.) had a preference for a soil moisture of about the wilting point
(12% ).
Observations in another year on a similar drought gradient (Ranga Rao et
al., 1991c) showed that Aphis craccivora was most abundant on plants that
were not drought stressed. This was in spite of being pounded with water
from an overhead irrigation system.
Wheatley et al. (1989) found that the the groundnut leaf miner was at an
advantage when its host was severely drought stressed. Extremes in precipi-
tation also appear not to influence the survivorship of the larvae of this species.
Shanower et al. (1992a) found no evidence that the extreme fluctuation
in population density that is characteristic of this species was in any
422 Groundnut pests
way related to rainfall events. Supplementary experiments showed that egg
and larva survival were not influenced by artificial rain for periods and
intensities that exceeded the normal range of field conditions.
11.3.3 Influence of insects on groundnl:d yield and economics

The development of an understanding of the relationships between the
feeding activity of insect populations and the yield of the host, with respect
to the characteristics of the farm system within which it is growing, is a
fundamental task facing all applied insect ecologists. In many cases, the
importance of natural enemies in the life system of a potential pest means
that the dynamics of three trophic levels have to be taken into consider-
ation (Shanower et al., 1992b). This implies the need for the construction
of predictive models. Despite the importance of such sets of information,
there are few reliable and published accounts relevant to pests of ground-
nut crops. This is unfortunate but is not surprising in view of the difficulty
of assembling and analysing the necessary data sets.
The relevance of this topic to insect pest management as a whole is that
these data are needed to establish parameters for the economic threshold -
the density (or intensity, in insect days) of an insect population at which its
activity results in the decline in the potential yield of the harvestable
component(s) (pods and/or haulm) of a crop.
The economic threshold can be distinguished from the action threshold,
which is the insect population density at which activity is needed to prevent
the population density exceeding the economic threshold at some time in
the future. Depending on the pest complex and the cropping system, the
'activity' could be:
• the commencement of hand picking or trapping the offending pest;

• the application of an insecticide;
• crop rotation; or
• deep ploughing, in the case of soil insect life systems.
A number of approaches have been adopted to secure relevant information

depending on the circumstances. The 'circumstances' are often in fact
governed by the inability of the entomologist to locate appropriate pest
populations.
(a) Energetics and laboratory feeding experiments

Most animals can be considered as machines that convert food (consump-
tion = C) into new body tissue (production = P), a process that exploits
part of the potential energy of the food to fuel the animal's metabolism
(respiration = R) of the food. Measurable by-products of this process also
include unassimilated ingesta (faeces = F) and excreta (U). The relation-
ship is:
C=P+R+F+U
In the last 20 years there have been many studies of this relationship,
especially with respect to the establishment of the trophic relationships of
phytophagous insects. They usually include estimates of the gross ecologi-
cal efficiency (PIC). This and other ecological efficiencies can be discussed
in terms of dry matter, energy and, more rarely, essential nutrients,
especially nitrogen, depending on the context of the study. For instance,
because energy supply is rarely limited in terrestrial systems, trophic
dynamics or system structure can be discusseCl in terms of the energy flow
from one trophic level to the next (Grimm, 1973; Axelsson et al., 1975;
Axelsson, 1977; Schroeder 1978; Wightman, 1979; Bellows et al., 1983).
Energy (or carbon) units, rather than units of mass, are adopted as the
common denominators when describing such systems because the 'concen-
tration' of energy - i.e. the number of joules per unit mass - is species and
possibly system specific. An extreme example is that of bruchid beetle
larvae living on dried pulse seeds. The cotyledon of the host, the larval
food, has an energy equivalent of 18.7 J/mg whereas bruchid larvae have
up to 27.8 J/mg (Wightman, 1978).
The constraints to energy flow or the 'bottle necks' in biological systems
are most likely to be revealed by studies of nutrient cycles. This is because
(after water-related problems) nutrient excesses or shortages are the most
common constraints on the components 0;: biological systems.
Measurements of the mass of system components are applicable to
studies where only changes over time are critical or where one component
is of primary importance. For instance, leaf mass consumed by an insect
can easily and accurately be considered in terms of the leaf area removed
(Schroeder, 1984) because this is directly related to the amount of light
energy intercepted by the host. Dry matter, or preferably ash-free dry
matter, is usually determined because variations in the water content of
most organisms (as influenced by environmental conditions) affect the
precision of biomass estimates.
Over the years it has become apparent that when a phytophagous insect
is feeding on a suitable host that is growing in stable conditions, the gross
ecological efficiency is in the region of 0.14 (mass) and 0.18 (energy)
(reviews by Edwards and Wightman, 1984; Wightman and Rogers, 1978).
This ratio will vary according to host species (and the genotype, if there are
variations in the level of allelochemicals within the species), the nutrient
status of the soil in which the host is growing and the degree of drought (or
water) stress (e.g. Mansour, 1981; Scriber, 1979a,b; Crawley, 1989). Thus
if the mass of the insect is known, it is possible to calculate the amount of
leaf material it has removed up to the time a measurement of mass
(or length, which is a function of mass) is made. It is then a matter of
424 Groundnut pests
TABLE 11.8 Leaflet area, fresh weight, dry weight and dry:fresh (dlf) ratio of 20
mature leaflets from five groundnut genotypes (Wightman, unpublished data)
Mean fresh wt drywt d/f mg (dry)

cm 2 g g cm- 2
Shulamit 199 4.88 1.49 0.305 7.48

VB 248 6.17 1.85 0.300 7.56
NC7 196 4.98 1.42 0.285 7.23
A46LlO 215 5.43 1.62 0.297 7.52
A 81 L 18 184 4.62 1.45 0.313 7.88
Mean 208.4 5.22 1.56 0.300 7.53
arithmetic to determine the effect of an insect population on the biomass or

leaf area index of a plant population or crop. This is the area in which plant
or crop growth models and insect models interface.
Estimation of larval damage

As an example, in the case of Spodoptera litura larvae feeding on ground-
nut foliage, their maximum length is c. 40 mm, which means that they
weigh 0.3 g dry (Rogers et al., 1976) which is equivalent to about 1.4 g live
weight). If PIC (mass) = 0.14, then C = 2.14 g.
Adopting the energy route gives a similar answer. Phytophagous insects
have an energy content of about 23 kJ/g (e.g. Edwards and Wightman,
1984; Schroeder 1977, 1978, 1984), thus P = 0.3 X 23 kJ = 6.9 kJ. If PIC
(energy) = 0.18, then C = 38.3 kJ. As leaves have c. 18.4 kJ/g energy
content (Petrusewicz and Macfadyen, 1970), this indicates that one larva
consumes 2.08 g of (dry) leaf during its development. As there are 7.53 mg
(dry) per cm 2 of groundnut leaf (Table 11.8) the mean (2.11 g) of these two
estimates is equivalent to 280 cm2 • Thus one larva consumes 20-30 leaflets
during its development, depending on the size of the leaf and proportional
mass of non-consumed vascular tissue.
Garner and Lynch (1981) measured the area of groundnut leaflet con-
sumed by S. frugiperda larvae (fall armyworm) in Georgia, USA. Their
main experiment showed that the mean leaf area consumed during the
larval period was 94.6 cm2 (which, following data in Table 11.8, is equival-
ent to 712.3 mg dry leaf). This is close to the cumulative consumption data
of ±100 cm 2 for the same species indicated by Smith and Barfield (1982).
The mean pupal mass was 177.5 mg. This means that the maximum
larval weight was about 213 mg, because pupae weigh c. 20% less than fully
grown larvae (Hagvar, 1975; Mackay, 1978; Wightman 1978). The dry
weight to live weight ratio of phytophagous insect larvae is normally about
0.2 (personal observation) so that the dry weight of the fully grown larva
was c. 43 mg. A PIC ratio of 0.14 indicates consumption of 304 mg, which is
considerably less than the observed.
Data from ancillary experiments carried out by Garner and Lynch (1981)
indicate that the disparity may be due to the age of the foliage with which
the larvae were fed. Adopting the data from follow-up experiments cer-
tainly brings the model and experimental data closer to agreement. Larvae
fed on 2-day-old leaves ate up to twice the area of leaflet, probably
bringing the PIC ratio closer to 14%. Furthermore, the larvae developed
more quickly, had a much lower mortality rate and finished somewhat
larger than counterparts fed on leaves up to 40 days old. This indicates that
antibiosis may have developed in the older leaves and was the cause of the
low estimated PIC ratio.
However, the possibility of a disparity between a model and the experi-
mental data calls for a re-examination of both. In this case we need to look
at several factors:
• Is the PIC really 6% for S. Jrugiperda or is this an artifact related to the
insects being fed old (and excised) leaflets?
• Are the Australian host data transferable to the US genotype?
• What are the equivalent experimental data for S. litura and other
Spodoptera spp.?
It is certainly an indication that model data should be applied with circums-
pection, and preferably with experimental verification. Huffman and Smith
(1979) present data indicating that Helicoverpa zea consumes 176 cm 2 of
foliage of the cv. Starr. However, we do not have access to biometric data
for this species.
(b) Simulation of defoliator damage in cage and field experiments

Continuing on the theme of conventional defoliators (which have attracted
most attention from experimentalists), several groups have evaluated the
effects of leaf removal by human or insect agencies on groundnut yield
(e.g. Greene and Gorbet, 1973; Enyi, 1975; Smith and Barfield, 1982). The
latter authors present a defoliation level (0-100%), by time (35-110 days
after emergence) and by yield reduction (0-50%) response surface for a
spanish variety. They conclude from this and other data: 'peanut is most
susceptible to defoliation from 70-80 days post emergence and practically
immune to yield reductions from defoliation prior to bloom initiation and
near harvest.' Certainly, the figure they present and the other data they
review support this conclusion.
At I CRISAT we released specially reared fourth instar Spodoptera litura
larvae onto plants that were surrounded by a 20 cm high metal barrier that
stopped their escape. This procedure was adopted to avoid the possibility
of delivering a systemic shock to the plants' system by abrupt hand or
mechanical ablation. In these experiments we have consistently found that
426 Groundnut pests
plants defoliated after the seedling stage or early flowering stage were
tolerant to attack. This was most marked in the rainy season. Plants even
produced pods (amounting to 50% of the control yield) when they were
completely defoliated from 10 days after emergence by the addition of two
larvae per plant at 20-day intervals (Wightman et al., 1990).
Similar experiments carried out by Sathorn Sirisingh and Manochai
Keerati-Kasikorn (1986) in Thailand indicated similar insensitivity to defo-
liation by groundnut plants.
The clue to the disparity in the results from North America and Asia
may lie in the genotype, their partitioning coefficients and sowing pattern.
In the experiments in India we expect canopy closure after 20-30 days. The
genotypes we experimented with (mainly spanish bunch) have a leaf area
index of >4 by the time the vegetative stage is complete. This means that,
in practical terms, after the crop has been in the ground for more than 30
days it can suffer at least 50% loss of leaf area before its photosynthetic
capability is reduced. This statement ignores the photosynthetic capacity of
the stems and petioles, which may be more than anticipated.
The principle behind cage experiments is to isolate plants from as many
insects and natural enemies as possible and then infest them with members
of the required species at whatever range of stage, time and density is
required. Primary assumptions are that the introduced insects, whether
reared or collected from another site, are healthy and are otherwise
'normal', as are the plants grown in cages. This approach has proved
satisfactory for establishing the relationships between the densities of
Nezara viridula and Riptortus serripes on soybean yield (Brier and Rogers,
1991) and Helicoverpa armigera on chickpea yield (ICRISAT Legumes
Program, 1991b). However, the cage experiments with groundnut leaf
miner at ICRISAT are believed to have given non-representational results
because the fine cage netting needed to secure this species intercepted too
much light, thereby changing the physiology of the plants.
(c) Field experiments

Experimental approach
Experiments in Thailand carried out by Sathorn Sirisingh and Manochai
Keerati-Kasikorn (1986) indicate that the groundnut leaf miner is capable
of more drastic yield reductions than S. litura. Their data indicate that, in
the season under discussion, a heavy infestation (c. 40 per plant) in the first
or second generation (flowering stage) reduced the yield from a potential
1.28 t/ha (site 1) or 0.95 t/ha (site 2) to 0.63 and 0.65 t/ha. The data also
show that the control of fungal diseases increased the above potential yield
by 25-50%.
The determination of the relationship between the density of groundnut
leaf miner populations and groundnut yield has also been the objective of
experiments carried out on the ICRISAT farm for 15 seasons (two per
TABLE 11.9 Groundnut yield, groundnut leaf miner intensity and GLM
parasitism as influenced by insecticide application (IIr generation)
% parasitism
Treatment Stand yield 8 Apr 85 19 Apr 85 insect 'days'

(pods) tlha
Dimethoate (400 g) 1.78 3.1 0 76.9

(8 applications)
Dimethoate (200 g) 1.70 13.6 33.9 752.4
(3 applications)
Diflubenzuron (250 g) 1.43 20.5 42.5 1443.2
(3 applications)
Dichlorvos 1.58 16.3 50.0 742.8
(3 applications)
Control 1.15 23.0 61.0 1617.0
(no insecticides)
SE ± 0.06 ± 3.5 ± 10.4 ± 67.0
The number of insecticide applications refers to the whole season (110 days).
Insecticides were applied in 350 I water/ha.
Rates (g) are a.i.lha.
year). Although this species was present in most seasons, it only reached a
density that could be described as damaging on three occasions. The
essence of these experiments is to allow the population to build up to
beyond our concept of an economic threshold and then to apply insecti-
cides in such a way that we achieve a range of population intensities and
control efficiencies. We then relate pest intensity to pod yield. These
experiments cover about 1 ha and are conducted in such a way that there is
a minimum of season-to-season variation.
With this species there is the possibility of several combinations of events
because it has three or four generations per growing season. It can there-
fore appear (and disappear) at any density between emergence and harvest
and display considerable variations in density change during a season. We
are, however, accumulating data that will enable us to put together a
rational management outline (Table 11.9). They indicate that an insecti-
cide application (e.g. dimethoate at 200-350 g a.i.!ha) should be applied
when there are five or more new mines per plant during the seedling stage,
10 new mines at flowering and 15-20 mines per plant up to 2 weeks before
harvest, after which insecticide application will have little impact on yield.
lassids and thrips often occur concurrently. Even if no other pest is
present, this makes it difficult to determine whether there is a relationship
between their intensity and pod or haulm yield. However, in the 1992 rainy
season at ICRISAT there was heavy infestation of jassids with minimal
thrips densities. The yield data (Table 11.10) indicate that there was a high
428 Groundnut pests
TABLE 11.10 Cumulative effect of jassid (Empoasca kerri) damage (eight days
before harvest) on haulm and pod yield of susceptible groundnut variety ICGS 44
(post-rainy season 1991/92, ICRISAT Center)
Insecticide Leaflets with Dry haulm Dry pod

applications jassid damage mass mass
(n)
% t/ha t/ha
0 60.0 3.26 1.73

1 31.3 3.20 1.91
4 17.0 3.55 2.14
SE ± 2.7 ± 0.22 ± 0.08
level of leaf damage by 8 days before harvest in unsprayed plots. However,

the reduction in the number of damaged leaflets by means of four insecti-
cide applications had no effect on haulm yield and little effect on pod yield.
Indications of the effects of soil insects on crop yields can be deduced
from the results of experiments involving the application of insecticides to
the soil. For instance, data provided by Kumawat and Yadava (1990)
indicate a linear relationship between the density of larval Holotrichia
consanguinea (white grubs) and plant mortality in experimental conditions
in Jaipur, Rajasthan, northem India. There was a log-log relationship
between density and pod yield.
Wightman et al. (1994) simulated white grub attack by cutting through
the roots of groundnut plants (White Spanish) 30 or 51 days after emerg-
ence (in glasshouse conditions) (Table 11.11). The root systems regrew
when the plants were cut after 30 days, although there was a considerable
energy cost in terms of reduced pod yield. Plants cut at the later date did
not regrow their roots. This was even more accentuated when the plants
were drought stressed. The plants with roots cut 51 days after emergence
were close to death at the end of this experiment.
This experiment demonstrated differential debilitation as a result of root
damage from flowering to harvest. But seedlings can also be killed when
attacked by white grubs (Bakhetia, 1982; Kumawat and Yadava, 1990).
Apart from the loss in yield, the farmers' profits are further reduced
because weeds are able to grow in the gaps. This reduces the opportunity
for compensatory growth by the plants next to the spaces left by the killed
seedlings. The potential role of compensatory growth following stand
thinning may, in any case, bc overestimated.
The extent and cost of seedling mortality and its management in north-
ern India have been estimated by Bakhetia (1982). He demonstrated
1.3-2.6% plant mortality where seeds had been dressed with insecticides
compared with 10.1% mortality where seeds were untreated. The overall
yield was at least doubled by using insecticide seed dressings, indicating
TABLE 11.11 Mean weight (n = 5) of pods produced when groundnut plants
(variety White Spanish) were cut through the root at 10,15 or 20cm below the soil
surface (0 cm = uncut control), 30 or 51 days after emergence when grown under
drought stress or fully irrigated conditions
Cut Mean pod weight g/plant ± SE
Drought stress No drought stress
Cut 30 days after emergence

Depth of cut
Oem 12.92 ± 1.64 22.96 ± 0.86
lOem 9.81 ± 0.14 14.19 ± 0.57
15 em 11.17 ± 0.36 15.11 ± 1.10
20 em 12.02 ± 0.09 15.91 ± 0.56
Cut 51 days after emergence

Depth of cut
Oem 12.92 ± 1.64 22.96 ± 0.86
lOem 7.73 ± 0.73 10.98 ± 1.08
15 em 9.36 ± 1.52 13.93 ± 0.47
20 em 8.99 ± 0.18 14.18 ± 1.50
that there was considerable additional sublethal yield loss that was avoided
by introducing insecticides into the soil as a seed coating.
Gough and Brown (1988) indicated that groundnut crops in the Atherton
Tablelands of north-east Australia were equally sensitive to attack by white
grub (Lepidiota sp.) Their data indicate that one larva per metre row (6-8
plants) of cv Virginia Bunch reduced crop yield by 381 kg/ha.
Within-stand compensation following plant mortality

Wightman and Wightman (1987) found that, in conditions typical of farming
systems in Malawi, there was no within-row compensation by plants in
stands that suffered up to 50% mortality once the stand had been above
ground for 26 days. ('Compensation' is defined as an increase in the pod
and/or haulm yield of plants in depleted stands, relative to plants in control
stands that are not depleted.) Up to this time there was within-stand
compensation for plant death only if there was> 30% mortality after 17 days
and >50% mortality at 26 days. These data refer to a crop that was harvested
after 5 months. They indicate, at least in the conditions of this experiment,
that compensatory growth of the pods and haulms in response to the death of
neighbouring plants occurs only in earliest stages of stand development.
The empirical or trial-and-error approach

Because the end point of the process under discussion is the develop-
ment of pest-management methods that are appropriate for the given
430 Groundnut pests
system, it is feasible to derive action thresholds by trial and error. For
instance, in Sao Paulo State, Brazil, where thrips are the major pest and a
serious constraint, farmers are advised to apply a suitable insecticide only
when there are more than three thrips per leaflet on 20% of a random 200
leaflets taken from 1 ha. This procedure reduced insecticide applications
from seven to two per season with an increase in yield and profitability
(Snhr Dalmo Lasca, Director, Sao Paulo State Extension Service, personal
communication). There is no experimental data to support this procedure,
but it works.
Similarly, at ICRISAT, we set up in 1984-85 a series of ad hoc action
thresholds to assist the farm manager's plant protection team before we
had supportive data. For instance, we recommended that an insecticide
(normally dimethoate at 200-350 g a.i.lha) should be applied for ground-
nut leaf miner control if the density exceeds five mines per seedling, 10
mines per plant at the flowering stage and 15-20 mines per plant up to 3
weeks before harvest, after which insecticide application is likely to have
little benefit. Our experimental data (above) indicate that this rule of
thumb had some merit.
11.3.4 Dynamic programming as a tool to guide research orientation

The action thresholds for the groundnut leaf miner just mentioned were
the basis of a series of modelling exercises carried out to evaluate manage-
ment scenarios for this pest. The exercise was based on a population
dynamics model of this insect and contemporary village-level fixed and
variable costs for southern India (Dudley et al., 1989).
The scenarios covered issues such as: 'If a farmer has available varieties
with 0, 10,20 ... 90% host plant resistance, how much natural mortality is
required at each level of host plant resistance to eliminate the need for
insecticide application?' The role of the market value in determining the
optimal number of sprays was also investigated. The final conclusions from
this piece of work are realistic and pointed to a difference between pest
management in developed and less developed countries. They point to our
need to make assessments of the effectiveness of farmers' insecticide
application activities and, implicitly, the role and effectiveness of natural
enemies (assumed to be inversely related to insecticide application activity)
before it is possible to work out what level of host plant resistance to a
given insect pest is needed.
11.4 HOST PLANT RESISTANCE
Host plant resistance, where it exists, can be made available to farmers as

an effective and environmentally friendly component of pest management
that involves little or no extra cost or effort than the normal purchasing,
sowing and keeping of seed - 'technology in the seed'. It is thus part of the
Host plant resistance 431
applied entomologist's job to detect and exploit it where feasible and
rational.
As inferred, the cost to the farmer is small; however, the institutional
costs are considerable and include the development and support of a
germplasm coIIection, research farm and research facilities, many years of
screening and selecting germplasm and probably about 10 seasons of
resistance screening and selection of breeders' material. This is followed by
on-farm testing and evaluation by farmers.
Of course, there is no guarantee that the traits sought exist in the
germplasm of the crop species, its close relatives or, in the context of
contemporary biotechnological feasibilities, any other species. There is
also no guarantee that the breeders and other gene-shifters will be able to
ensure that the desirable genes to manifest themselves in a variety that is
adapted to the target environment. Thus, breeding for host plant resistance
is primarily an activity of organizations that are stable, mission-orientated,
well endowed and non-profit-making.
Many groundnut genotypes have characteristics that protect them from
herbivores. This observation is based on the relatively small number of
pests (as opposed to insects) associated with the above-ground parts of this
species. (We have yet to come to terms with resistance to root-eating
arthropods.) Lynch (1990) has made a notable contribution to groundnut
science by listing the genotypes that are known to have resistance to many
of the most important pests.
If a particular insect is established as being a major constraint in one or
several agroecological zones, it is rational to consider the inclusion of host
plant resistance in a groundnut management programme that includes the
provision for breeding or selecting adapted varieties. If no source of
resistance genes is known, it is necessary first to develop guaranteed
screening procedures for distinguishing between resistant, susceptible and
'escape' plants or genotypes. Rationally, this process should also enable
other characteristics of the screened genotypes to be assessed. Screening
methods are not covered here but the general principles are described by
Smith (1989).
Once resistance to one or several potential pests has been found, it is
usually necessary to work with breeders to combine the relevant genes with
a 'background' that is agronomically suited to the target environment. This
procedure is made more efficient if it is possible to supplement the field
screening of progeny by monitoring the presence or absence of the physical
or chemical markers associated with the mechanism of resistance (Lund-
gren et at., 1981, 1982).
11.4.1 What is host plant resistance?

Host plant resistance is a phenomenon that has evolved in most higher
plants to permit them to coexist with or to avoid the many species of
432 Groundnut pests
potential herbivores that could exploit them as food. Viewed broadly, it
can take several forms that are usually connected with the feeding activity
of the free-living forms of the herbivores or the provision of food and
shelter for their progeny.
1. Repellence (antixenosis, 'non-preference' or the turning away) of herbi-
vores before they come into contact with the plant. This can be associ-
ated with, for instance, the release by the plant of physiologically active
chemicals (kairomones) into its air space or with a physical factor that
influences the herbivores' visual response to a plant (or group of
plants), perhaps to the extent that it is not recognized as a potential
host. Physical characteristics such as the presence or absence of tri-
chomes on leaf or stem can also influence the way that an insect reacts
to a plant when first approaching it.
2. Antibiosis, where the plant contains chemicals that, when tasted or
ingested by a herbivore, prove to be antimetabolites (e.g. insect growth
hormone analogues), repellents, antifeedants, or toxins (including the
toxic manifestations of the genes from other organisms, such as Bacillus
thuringiensis) .
(a) Latent antibiosis awaits being switched on by a challenge from a
herbivore or by a systemic (within plant) or pheromonal (between
plant) message. Latent resistance has not (yet) been detected in
groundnut although it may exist; it is a factor in the resistance of
tomato plants to Spodoptera littoralis (Edwards et al., 1985).
Further information about this phenomenon can be found in Kogan
(1986) and Edwards and Wratten (1987).
(b) Temporary antibiosis is only present during a particular stage in the
development of a given organ or the phenology of a plant.
(c) Permanent antibiosis is a characteristic of a given plant species or
organ.
Antibiosis can also take the form of the absence or masking of a feeding
stimulant.
3. Tolerance, where the plant can continue to develop and reproduce
despite being attacked by herbivores. The misuse of this term to denote
low levels of antibiosis or an undefined aspect of the resistance phenom-
enon, in general, often leads to unnecessary confusion.
4. Physical (structural), where the plant has structures (trichomes, thorns)
or surface characteristics (thick or waxy cuticle, or even a layer of
water - Nwanze et al., 1990) that interfere with a herbivore's ability to
exploit it.
This is a development of the conventional view of host plant resistance in
plants to herbivores, based on Painter (1951). The following could be
added because they can be complementary or confounding:
5. Avoidance (seasonal), where the plant's phenology (or a crop's sowing
pattern) is such that its life-cycle (or a sensitive developmental stage
such as flowering or seed swelling) does not coincide with the time of
year when a key herbivore is active.
6. Avoidance (spatial), where a plant has evolved into forms that grow
(or are sown) outside the biogeographic or climatic range of a key
herbivore.
However, in practical terms, avoidance is best considered as being an
aspect of cultural control.
The genus Arachis displays several of these resistance factors.
11.4.2 Host plant resistance in Arachis hypogaea
(a) Resistance to Aphis craccivora

Although seedling pests in their own right, groundnut aphids have been
highlighted as primary groundnut pests in Africa because of their ability to
transmit the groundnut rosette virus complex (GRV). In one season (1975)
they converted Nigeria from one of the world's leading groundnut export-
ing nations to relative obscurity in this regard.
Considerable energy and expertise has been devoted to breeding for
GRV resistance per se but it is surprising (with the benefit of 40 years of
hindsight) that the colonial authorities of the time did not follow up the
discovery made by Evans (1954) in Tanzania. He found that several
varieties, especially Asiriya Mwitunde (a name indicating that the variety
belongs to the Mwitunde people of northern Tanzania), carried compara-
tively small aphid popUlations compared with the other varieties tested.
This was associated with fewer and smaller GRV primary infestation sites.
This clear indication of virus management by vector control was accomp-
anied by significant yield advantages.
Unfortunately, screening of the East African genotypes in the ICRISAT
germ plasm collection has only revealed comparatively low levels of aphid
resistance. Apart from the implication that important germplasm has not
yet been collected or has been lost, this is now of less importance because
two genotypes with high levels of aphid resistance, ICG 5240 (= EC 36892)
and ICG 5725, have been detected (Wightman et ai., 1990). Field tests of
ICG 5240 in Africa (Wightman et ai., 1990; Sithanantham et ai., 1991),
India (Padgham et ai., 1990a,b) and China (Dr Xu Zeyong, personal
communication) have shown that the level of aphid resistance remains high
across continents. These experiments have shown sufficient intercontinen-
tal variation in aphid response to demonstrate the existence of biotypes of
A. craccivora. Resistance is manifested by longer generation time and
considerably diminished fecundity.
Field tests in the rigorous conditions of a GRV screening nursery in
Malawi showed that after 40 days exposure (after which GRV has a minor
434 Groundnut pests
effect on yield) 14% of the 248 ICG 5240 plants were infected (control =
70% infestation, n = 749). At harvest time, 4 months after emergence, the
susceptibles had a >98% infestation, and ICG 5240 only 44%.
The next step in exploiting this resistance was to determine the mechan-
ism. This has given us access to a method for screening the progeny of
crosses between the aphid resistant genotypes and lines with agroecological
adaptation that would complement or replace our bioassay technique.
These studies (carried out mainly at ICRISAT by scientists from the
Natural Resources Institute, Chatham, UK) have shown that there is only
evidence for antibiosis. It is temporary in that the factor is concentrated in
the areas where the aphid is most likely to feed, i.e. the terminal leaf buds,
leaflets and petioles. Electronic monitoring showed that the aphids fed for
half the time on the phloem of ICG 5240 compared with TMV 2 (suscep-
tible control). This is likely to reduce considerably the chances of absorp-
tion of the GRV virus by the vector (now known to be a complex of
viruses, e.g. Murant, 1990; Murant and Kumar, 1990), which has an
acquisition time of >4.5 h (Padgham et al., 1990a; Padgham et al., 1990b).
The analysis of phloem extracts from the petioles of ICG 5240 had 2-8
times the concentration of procyanidin than did the phloem of TMV 2.
Further tests showed that there was a strong negative correlation (r2 =
-0.86) between the log procyanidin concentration in seven groundnut
genotypes and the intrinsic rate of increase of aphids on those genotypes.
Assays showed that a concentration of only 0.005% procyanidin in an
artificial diet reduced honeydew production (equivalent to diet ingestion)
by 50% (Grayer et ai., 1992; Kimmins personal communication). This
implicates procyanadin in the resistance process, though not necessarily as
the active component: it is, however, a convenient quantitative marker
that is being used as an index of resistance level.
Thus resistance to the vector of GRV has been detected and the possible
mechanism of resistance described. Vector resistance has been shown to be
a valid method of managing the disease and there is a relatively simple
method of detecting minute quantities of the resistance factor (or a precur-
sor or breakdown product) in plant sap.
(b) Resistance to thrips and jassids

Lynch (1990) and Wightman et ai. (1990) list more than 100 genotypes with
resistance to thrips and/or jassids. Subsequent screening at ICRISAT has
revealed several more - not detailed here, except that it is worth mention-
ing that ICG 5240 has high resistance to jassids as well as to aphids (above)
and foliar diseases (Sithanantham et ai., 1990). However, at this stage in
the development of pest management procedures for groundnut, it is more
appropriate to develop varieties with high yield potential that also have
resistance to key pests and groups of pests than to seek more resistant
genotypes when plenty have already been identified. Table 11.12 indicates
TABLE 11.12 The pedigree, yield and jassid score of 10 out of 18 groundnut
varieties bred for jassid resistance and high yield, tested in the 1990 rainy season at
ICR1SAT Center (communicated by Dr S. L. Dwivedi)
Variety Pedigree Pod yield Jassid

(ICGV) tlha score*
87745 (SB) ICG 799 x (ICG 799 x NC Ac 2214) 2.54 3.3

86455 (SB) ICG 799 x (ICG 156 x NC Ac 2214) 2.40 3.0
86393 (VB) ICG 1326 x (ICG 156 x NC Ac 2214) 2.48 3.0
86462 (SB) ICGS 1 x NC Ac 2240 2.31 2.3
86522 (SB) ICG 799 x (M 13 x NC Ac 2214) 2.20 3.0
86518 (SB) ICG(PRS) 12 x NC Ac 2214 2.20 2.3
87430 (SB) ICGS 6 x (ICG 799 x NC Ac 2214) 2.16 3.0
87252 (SB) ICGS 7 x NC Ac 2214 2.15 2.0
87495 (SB) F334A-B-14 x NC Ac 2214 2.12 3.0
87468 (SB) ICGS 24 x NC Ac 2214 2.04 2.7
Controls
High yielding susceptible:
ICGV 87128 (SB) 2.36 7.0
Resistant parents
NC Ac 2214 (lCG 5040) (VR) 1.51 1.0
NC Ac 2240 (ICG 5043) (VR) 0.39 1.0
Trial mean (total of 18 entries) 2.25 3.5
SE ± 0.17 0.2
Notes: SB = spanish bunch. VB = virginia bunch. VR = virginia runner;

*Jassid score 1 = no damage (chlorosis). 9 = >75% of leaves are chlorotic;
Data in parentheses are the ranking of yields within the experiment.
that this procedure has led to the combination of jassid resistance genes
from ICG 5040 (low yield potential) and ICG 5043 (very low yield poten-
tial) with backgrounds conferring relatively high yield potential in research
station conditions (Dr S.L. Dwivedi, Groundnut Breeder, ICRISAT).
They now have to be tested in farmers' fields in appropriate areas.
Screening for resistance to thrips and jassids in Thailand over a number
of years has indicated that (NC Ac 343 x NC 17367) had the highest yields
of the lines tested and multiple insect resistance. NC Ac 343, (NC 1107 x
(NC 2232 x NC 2214» and (NC 6 x NC 3033) had sightly lower yields but
high levels of pest resistance (investigators were W.V. Campbell,
Manochai Keerati-Kasikorn and Turnjit Satayavirut; the latter communi-
cated this information). This agrees with findings previously reported for
resistance to Frankliniella fusca in North Carolina in which NC Ac 343 and
its derivative NC 6 were linked with thrips resistance (Campbell and
Wynne, 1980).
436 Groundnut pests
TABLE 11.13 Incidence of Thrips palmi and bud necrosis disease (BND) in an
experimental trial carried out at Rajendranagar, near Hyderabad, 1992 post-rainy
season (means of four replicates, data of Vijaya Lakshmi (unpublished); JI 24 =
susceptible control)
Days after Genotype Thrips palmi BND incidence

sowing n125 terminal % of all plants
29 ICGV 86031 9.5 0

ICGV 86338 9.8 0
JI24 29.8 0
44 ICGV 86031 9.8 0
ICGV 86338 13.8 0
JI24 14.8 1.2
57 ICGV 86031 4.5 0
ICGV 86338 4.3 0.6
JI24 5.3 3.6
70 ICGV 86031 1.5 0
ICGV 86338 1.8 3.6
JI24 6.0 8.0
85 ICGV 86031 0.5 0
ICGV 86338 0.5 3.6
JI24 l.0 18.7
99 ICGV 86031 0 0.6
ICGV 86338 0 3.6
JI24 3.8 24.8
114 ICGV 86031 0 0.6
ICGV 86338 1.3 3.6
JI24 2.0 33.6
132 ICGV 86031 0.3 1.2
ICGV 86338 0 4.2
JI24 0 39.1
Resistance to the thrips vector (Thrips palmi) of the bud necrosis virus
(BNV - a variant of the tomato spotted wilt virus (TSWV) that is found in
the Indian sub-continent) is viewed as being an important key to the
management of this disease in groundnut (Amin, 1985a; Reddy and
Wightman, 1988; Reddy et al., 1991). A variety of great potential in this
respect is ICG 86031 which has resistance to the vector and to other insects
(below) and has unconfirmed resistance to the virus. Field trials have
indicated that ICGV 86388 also has resistance to the vector which results in
low levels of BNV incidence (Table 11.13). In the USA, Southern Runner
suffers less from TSWV than Florunner (Culbreath et al., 1992).
High trichome density, distribution and length have been shown to be
important resistance factors in genotypes such as ICG 5040 (NC Ac 2214)
and ICG 5043 (NC Ac 2240) (CampbeJl et al., 1976; Dwivedi et al., 1986).
However, the important line ICG 2271 (NC Ac 343) and its derivatives are
not particularly hairy, so that presumably there is a chemical basis to its
antibiosis. As in the wild species (below), a flavone glucoside has been
linked with antibiosis in the cultivated species (Holley et al., 1984).
(c) Resistance to the lesser cornstalk borer (LCB)

In view of the importance of this insect to the peanut industry in the USA,
it is not surprising that resistance to this species has been sought on several
occasions. Smith et al. (1980a,b) screened 490 accessions in artificial con-
ditions and indicated that varieties Early Runner, Florigiant, Florunner
and Virginia Bunch were among the resistant lines. Field tests in North
Carolina of an initial 120 lines with natural infestations (i.e. screening was
carried out in realisitic conditions) were carried out from 1976 to 1981. PI
269116 ranked first or second for lowest peg and pod damage in the four
seasons in which infestations were heavy enough to give good screening
conditions (Stalker et al., 1984). Tests carried out in North Carolina
indicated that several lines are promising.
(d) Resistance to Spodoptera litura

The development of resistance to S. litura in suitable varieties has been
regarded as being of high priority for Asian groundnut farmers for a
number of years. The results of experiments carried out in 1986 and 1987
(Table 11.14) indicated the possibility that ICGV 86031 (breeder = S.L.
Dwivedi, ICRISAT) had some resistance to S. litura combined with high
yield in the post-rainy season. This hope was substantiated in further tests
on the ICRISAT research farm and in farmers' fields in coastal Andhra
Pradesh (southern India). In the limited trials so far carried out, farmers
had sufficient confidence to grow this variety without protecting it with
insecticides. They were rewarded with higher yields and lower variable
costs than neighbours who grew locally acceptable varieties but applied
insecticides to kill defoliators. PI 269116, PI 269118 and PI 262042 had
resistance to this insect but none were outstanding (Campbell and Wynne,
1980).
Bioassays carried out with larvae as preliminaries to detecting the mech-
anism of resistance (independent tests by Ranga Rao and Dr D.E.
Padgham, NRI) revealed no antibiosis effect on II-VI instar larvae when
fed mature leaves of ICGV 86031. The main mechanism of resistance is
currently thought to be tolerance, manifested as the enhanced ability of
vegetative tissue to regrow following defoliation.
However, first instar larvae suffered 56% mortality when fed on ICGV
86031 compared with 12% mortality when fed on susceptible ICG 221.
Padgham also found that newly hatched larvae had a marked propensity to
vacate the leaves of this variety in the first two hours of free life. This
TABLE 11.14 Effect of releasing two fourth instar Spodoptera litura larvae on the percentage defoliation (%d) (assessed after the larvae
had pupated) and subsequent mean plant yield (y) in g per plant of five groundnut genotypes (the data are the means of five replicates;
percentages were transformed to arc-sines for ANOVA)
Genotype Days after emergence
10 30 50 70 Control
%d y %d y %d y %d y %d y
Post-rainy season
ICGV 86031 86 10.0 58 11.7 37 15.0 22 18.8 18.3
ICG 5240 61 8.4 70 8.3 44 6.9 11 14.5 14.5
ICGV 86535 83 10.2 68 11.7 54 10.4 12 12.8 12.5
ICG 156 100 6.1 75 13.3 41 13.1 28 17.1 17.9
ICG 221 100 4.4 91 7.9 50 10.0 18 12.8 13.7
SE for: %d = ±1.6 (calculated via arc-sine transformation)
g =±1.3
Rainy season
ICGV 86031 100 4.5 45 9.2 64 10.8 35 10.8 11.4
ICG 5240 100 10.5 54 10.0 67 13.3 24 12.0 14.7
ICGV 86535 100 3.9 47 9.2 64 10.9 26 9.6 12.0
ICG 156 100 4.7 44 10.0 60 10.2 38 12.5 10.4
ICG 221 100 1.7 50 4.0 69 5.3 39 5.4 5.4
SE for: %d = ±0.5 (calculated via arc-sine transformation)
g = ±1.1
10 days after emergence (DAE) = seedling stage, 30 DAE = flowering, 50 DAE = pegging, 70 DAE = pod filling. Control plants had no insects and were
insect-free throughout the experiment.
suggests that the resistance factor that influences the neonates is associated
with the leaf surface, because their feeding activity is restricted to scraping
the leaf surface. The antixenosis demonstrated by ICGY 86031 is likely
to increase the first instar mortality that is characteristic of r-strategist
noctuids (Kyi et al., 1991) and will therefore contribute to the determi-
nation of the level of damage caused by the older larvae, among which
mortality is comparatively low.
(e) Resistance to the corn earworm (CEW)

Campbell and Wynne (1980) report resistance to the CEW in Early Bunch
and NC 6. The resistance in NC 6 affects larval development and is most
likely to be indicative of antibiosis.
(f) Resistance to the southern corn rootworm (SCR)

Experiments in North Carolina carried out over nearly 20 years resulted in
the development of NC 6 (= NC Ac 343 x Y A 61R) which, with other NC
Ac 343 crosses, competed favourably with Florigiant in terms of quality
and price. It also had 10-20% of the damage caused by SCR and did not
need protection from this pest under high infestation pressure (Campbell
and Wynne, 1980).
(g) Resistance to the groundnut leaf miner

Resistance to the groundnut leaf miner has been as difficult to locate. This
is mainly because of the sporadic nature of the infestations and difficulties
in performing realistic screens on a large number of genotypes with artifici-
ally reared insects. The difficulties include the need to exclude the parasites
but to allow plants to grow in a natural light regime without shading. The
inference is that fine net cages (needed to exclude parasites) cut down the
amount of light reaching the plants, and that supplementary artificial light
appears to cause variations in the leaf chemistry that modify resistance
factors (P.l. Moss, unpublished MSc thesis). An indoor screening process
that gives a satisfactory degree of consistency has been developed.
Progress has also been made with contributions by P.W. Amin, Dr R.Y.
Satyanarayana Rao, (Indian Agricultural Research Institute) and Ms P.l.
Moss (University of Bath, UK) in that resistance (tolerance, antibiosis and
physical) to this species has been detected. Tests have reached the stage
where trials in farmers' fields are called for to evaluate the significance of
this resistance in relevant conditions. Noteworthy genotypes are ICG 2271
(NC Ac 343), ICG 1697 (NC Ac 17090) and ICG(FDRS) 4. Anderson et al.
(1990) found that NC Ac 2821 (as well as NC Ac 17090 and PI 405132) had
resistance to the groundnut leaf miner when tested in Khon Kaen,
Thailand.
440 Groundnut pests
In contrast to jassids and thrips, it appears that groundnut leaf miner
moths are attracted to hairy leaves, as opposed to shiny (glabrous) ones for
oviposition (R.V. Satyanarayana Rao and G.V. Ranga Rao, unpublished).
Females are attracted to NC Ac 2214 (lCG 5040) for oviposition but the
larvae that hatch subsequently suffer higher levels of mortality because of
the antibiotic properties of this genotype. The latter appear to be associ-
ated with the exudation of relatively large volumes of sap when the plant is
injured. The sap is 'gummy' , inferring the possibility that the activity of the
larvae may be impaired, but it also contains comparatively high concen-
trations of polyphenols which have been associated with resistance to
Spodoptera litura in groundnut.
(h) Resistance to soil insects

Amin et al. (1985) reported that several of the lines that have resistance to
jassids and thrips also suffered less from termite scarification than the other
genotypes tested. Subsequent testing in Burkina Faso confirmed this
finding in field conditions (Lynch et al., 1986). Important lines include ICG
2271, ICG 5043, ICG 5044 and ICG 5045 (= NC Ac 343, NC Ac 2240, NC
Ac 2242 and NC Ac 2243).
Resistance to white grubs has not yet been located in Arachis. Trials
carried out with the above termite resistant lines in Australia, testing for
resistance to the pod feeding Heteronyx, revealed no resistance (H. Brier,
personal communication). However, resistance to white grubs has been
found in other crops. Crocker et al. (1990 ) recorded considerable levels of
resistance to Phyllophaga congrua in one wheat and four oat cultivars.
Lucerne, lupins and Lotus are highly resistant to Costelytra zealandica
(Farrell and Sweeney, 1974; Kain and Atkinson, 1970). Lucerne has been
sown in New Zealand as a cleansing crop in dry, lowland pastures that have
been badly affected by this pest since the beginning of this century.
Lucerne is also resistant to Heteronyx arator, another white grub pest of
pasture in New Zealand and Australia (King et al., 1975).
11.4.3 Host plant resistance in Arachis spp.

Many Arachis spp., the 'wild species', have levels of insect resistance that
approach immunity, i.e. insects walk away from them or, if confined on the
leaves, die if they eat them or die of starvation rather than eat them. This
phenomenon has been known for many years (Smith and Barfield, 1982;
Amin, 1985b) but, unfortunately, the exploitation of this knowledge has
been insufficient in view of the economic importance of some of the insect
species involved.
The extent of resistance in the wild species is shown in Table 11.15 which
summarizes the results of tests carried out at ICRISAT Center between
Natural enemies of groundnut insects 441
1988 and 1992. This represents a formidable amount of potentially exploi-
table material, especially when it is realized that the levels of resistance far
exceed that detected in A. hypogaea to the more intractable insects. The
inclusion of a root feeding species (jewel beetle) is an indication that a
search for resistance to the soil insect pests of groundnut among the wild
species may be justified.
It is also noteworthy that 6 of 18 (A. hypogaea x A. cardenasii) inter-
specific derivatives were resistant to the groundnut leaf miner in research
station (open field) conditions in Tamil Nadu, India (Kalaimani et al.,
1989).
However, progress has been made in other directions in that a start has
been made in determining the mechanisms of resistance in 14 species
(Stevenson et al., 1993a, b; Kimmins et al., 1993). Bioassays (Table 11.16)
indicated that the survival and growth rates were, in all cases, significantly
lower than in the susceptible A. hypogaea control (leG 221 = TMV 2).
Estimates of leaf toughness (,biteability') indicated that physical factors
may be components of this resistance phenomenon.
Bioassays of solvent extracts of the leaves of the most resistant species,
A. paraguariensis and A. chacoensis and an F} hybrid (A. chacoensis'X A.
hypogaea) indicated the presence of biologically active (antibiotic) frac-
tions which were quercetin glycosides that resembled chlorogenic acid.
Subsequent tests have indicated the flavonoid diglycosides that are present
in the leaves and may be the main resistance factors (Table 11.17).
Thus resistance has been found within the genus to reduce the effects of
the serious above-ground pests in groundnut. In several cases, there is no
scientific reason why this phenomenon should not be helping farmers in a
number of countries. The gaps in our knowledge point to the need to find
resistance to soil insects and to exploit the wild species.
11.5 NATURAL ENEMIES OF GROUNDNUT INSECTS
Not much is known about the dynamics of the natural enemies of ground-
nut pests. There are exceptions, but they are from the southern states of
the USA and emanate from institutions which provide strong linkages
between high quality research organizations and extension services that are
well tuned to the needs of local farming systems. Elsewhere in the world
we are somewhere between a zero knowledge base and the stage where
research reveals the need to carry out more research rather than the
solutions to problems.
11.5.1 North America

Smith and Barfield (1982) and Lynch and Douce (1992) are positive about
the potential for the natural control of Heliothis zea (corn earworm) and
442 Groundnut pests
TABLE 11.15 Arachis spp. tested at fCRfSA T Center for resistance to Aphis
craccivora (screen house), groundut leaf miner (GLM) (field), jewel beetle larvae
(field), and Spodoptera litura (laboratory assay) with an indication of the level of
resistance
Collection Arachis Groundnut Groundnut Jewel Army

number species aphid leaf miner beetle worm
10002 apressipila HR S HR
9990 apressipila HR R R
9993 apressipila HR S HR
30003 apressipila HR R HR
30009 apressipila HR R HR
30080 batizocoi HR S
30079 batizocoi HR
30081 batizocoi HR HR
GKP 9667(316) batizogaea HR
36034YO-l cardenasii HR S S
36034YF cardenasii HR R S
36019-1 cardenasii R
36033Y cardenasii HR R
GKP 10017 cardenasii HR
10602 chacoensis S S R
10602-5 chacoensis R
10602(5) chacoensis HR
30109 paraguariensis R
565-6 paraguariensis S
HLKHe565-6 paraguariensis HR
30134 paraguariensis HR HR
KCF11462 paraguariensis R
30109 paraguariensis HR
GKPSC30124 paraguariensis HR
9634 pseudovillosa R
12922 pusilla HR R HR
114 rigonii S
30007 Arachis sp. R HR R
A77/113 Arachis sp. HR
GKP9578(312) Arachis sp. S
GKP9797 Arachis sp. HR
GKPSCS30135(21) Arachis sp. HR
GKP9893(321) Arachis sp. HR
9921(100) Arachis sp. HR
KG30012(339) Arachis sp. HR
GKPSC30116/19 Arachis sp. HR
GKPSC30114 Arachis sp. HR
TABLE 11.15 Cont.
Collection Arachis Groundnut Groundnut Jewel Army

number species aphid leaf miner beetle worm

GKPSCS30135 Arachis sp. HR
GKP 10602 chacoense IR
36025-1 chiquitana S HR
9530 correntina S R
K 7988 duranensis HR S S
30074 duranensis R
30065 duranensis S
30067 duranensis S
30070 duranensis HR
HL 189 glabrata HR
HLKHe571(9IB) glabrata HR
A45/114 hagenbeckii HR
A27/117 hagenbeckii HR
2A5 hagenbeckii HR
GK 30006 hoehnei R
30085 kemf-mercadoi R HR HR
35001 kemf-mercadoi HR
30035 khulaminii HR S HR
30063 monticola S
7264 monticola R
30008 otavoi R S
30017 otavoi HR S
GKPSBS30132(1) Arachis sp. HR
GKPSC30122 Arachis sp. HR
GKPSC30120(13) Arachis sp. HR
Manfredi-5 Arachis sp. HR
10038LL spegazzini S
30126 stenophylla HR HR HR
HLK410 stenosperma HR HR S
HLK 408 stenosperma HR HR S S
HLK 409 stenosperma HR R
30011 valida S R
villosulicarpa HR S
villosa HR
TMV2 hypogaea S S S S
EC 36892 hypogaea HR S S S
HR = highly resistant; R = resistant; S = susceptible

444 Groundnut pests
TABLE 11.16 Insects and mites to which resistance has been located among the
wild Arachis spp. and interspecific derivatives (details are in Lynch (1990) or in
Table 11.15
Thrips Campbell and Wynne (1980)

Amin (1985b)
Empoasca spp. Campbell and Wynne (1980)
Amin (1985b)
Tetranychus spp. Johnson et al. (1977)
Leuck and Hammons (1968)
Lesser corn stalk borer Kamal (1978)
Stalker et al. (1984)1
Groundnut leaf miner Table 11.15
Kalaimani et al. (1989)
Spodoptera spp. Lynch et al. (1981)
Kimmins et al. (1993)
Table 11.15
Heliothis zea Stalker and Campbell, (1983)
Southern corn rootworm Stalker and Campbell (1983)
Root feeders Table 11.15
Aphids Table 11.15
Amin (1985b)
lThe authors noted that cultivatcd gcnotypes were as resistant as the wild species tested and
suggested that there was no need to attempt to exploit the wild species for resistance to this
species.
TABLE 11.17 Flavonoid diglycosides in leaves of most resistant Arachis species
Species Flavonoid diglycosides Concentration

in fresh leaf
(mM/g)
A. chacoensis Quercetin 3-arabinosylgalactoside 3.10

Quercetin 3-digalactoside 1.50
A. chacoensis Total 2.50
x A. hypogaea
hybrid
A. paraguariensis Quercetin 3-rhamnosylgalactoside 1.80
Kaempferol 3-rhamnosylgalactoside 1.56
A. hypogaea Quercetin 3-digalactoside 0.46
(TMV2)
H. virescens (tobacco budworm) in the south-eastern USA. Egg parasites
(Trichogramma sp.) giving 3-83% mortality, larval parasites (Microplitis
croceipes and Eucelatoria armigera) and nuclear polyhedrosis virus com-
bine to maintain defoliator population densities below economic status.
Smith and Johnson (1989) undertook a 3-year study, covering six dis-
cernible generations, of the population dynamics of Elasmopalpus ligno-
sellus (Jesser cornstalk borer, LCB) in Comanche County, Texas, basing
themselves in fields of cv. Starr that had no insecticide applied, no irriga-
tion and a history of LCB attack. They found that within-generation
mortality ranged from 87.1 % to 96.5%. Although larval mortality could be
ascribed to an entomopox virus, a fungal disease, 13 primary parasite
species and five species of predator, the main (key) mortality factors were
'unidentified' and density independent, and they influenced the eggs
(average 7.4%) and first instar (average 53.8%). This is not an unusual
feature of the population dynamics of R-strategist Lepidoptera (Kyi et al.
1991).
The implication of density independent mortality factors as being more
important than parasites, predators and pathogens is consistent with the
observation by Lynch and Douce (1992) to the effect that parasites and
predators maintain LCB population densities at sub-pest levels except in
seasons that are abnormally dry and hot. The host larvae are able to
tolerate these conditions whereas the natural enemies are not. This results
in outbreaks of the species in conditions where the crops are potentially
least tolerant to additional stress. We now need to know the nature of the
unidentified mortality factors and how or if they are influenced by the
temperature and moisture content of the soil within the larval zone of
activity. The possibility that changes in nutrient status of the plants that are
associated with drought stress also favour the proliferation of this species
(Wheatley et al., 1989) has apparently not been considered.
The potential importance of spiders in groundnut fields is highlighted by
a very detailed study carried out in Texas on irrigated and dryland fields
that had not been treated with insecticides during the 1981 and 1982
seasons (Agnew and Smith, 1989). More than 25000 spiders were col-
lected, belonging to 18 families and 79 genera. Hunting spiders out-
numbered web spinners by about 10 to 1. The list of prey is interesting
because it includes a number of potential pests, such as mirids, larval
Heliothis sp., jassids, and thrips. However, of the 220 prey records, 72 are
of predacious insects (excluding Hymenoptera), including 38 spiders, and
21 are Hymenoptera (14 ants and 6 parasitica). This indicates that spiders
confounded ecologist's concepts by operating in two trophic levels.
Earwigs (Labidura riparia) predominated in a 2-year study of arthropod
predators in peanut fields in Alabama (Kharboutli and Mack, 1991). They
ate caterpillar larvae (LCB, CEW and FA W). The exponential increase
and decrease in their density was interpreted to be associated with the
rapid exploitation of their food source and its subsequent exhaustion. The
446 Groundnut pests
voracious and polyphagous red imported fire ant Solenopsis invicta was the
next most abundant predator.
The stable and efficient operation of such natural control systems within
agroecosystems is dependent on the presence or provision of off-season
refuges that support a reservoir of hosts and their food plants. Such
systems are also easily disrupted by insecticides to which, in our experi-
ence, spiders have a particularly low tolerance. We wonder to what extent
the status quo for Heliothis spp. of the early 1970s still exists in commercial
areas, and whether it could be improved upon by the exploitation of host
plant resistance.
The results of recent research carried out at ICRISAT make us approach
this notion with caution for two reasons. Firstly, it does not include
reference to the potential confounding effect of host plant genotype on the
rate of parasitism. Secondly, the fourth trophic level, which includes the
parasites of parasites (hyperparsites), can also influence the sUccess of
natural control processes.
R.E. Lynch and 1.1. Hamm (unpublished) demonstrated that, in the
southern USA, the nuclear polyhedrosis virus (NPV) of H. zea was
compatible in a tank mix with chlorothalonil applied for leaf spot control.
The viability of the virus was reduced by about 60% but this did not
prevent the initiation of epizootics in field trials. Subsequent population
densities of the target species were maintained at below the economic
damage level because natural enemies were not affected by the virus.
11.5.2 Asia
There is little doubt that, in insecticide-free conditions, natural control
processes can have a marked effect on the densities of the pan-Asian
defoliators. A list of the natural enemies of Spodoptera litura has been
compiled (Ranga Rao et al., 1993). It includes:
69 parasite species in 7 hymenopterous and 2 dipteran families
36 species of predacious insect in 14 families
12 species of spider in 6 families
4 species of protozoan
4 species of fungal pathogen
7 bacteria
4 viruses
5 nematode species
This probably represents the tip of the iceberg, because it represents what
happens when a suitably qualified person with the means to observe, rear,
collect, identify and report is in the right place.
The levels of larval and egg parasitism in S. fitura are monitored on the
ICRISAT farm (100-400 individuals or egg rafts) during each season. The
data indicate that the tachinid Paribaea orbata is the most common larval
Natural enemies of ground nut insects 447
parasite and that Ichneumon spp. and Exorista xanthopis also contribute to
larval mortality. The rate of larval parasitism is 9.0% over eight seasons,
but this includes two seasons where it reached 26.2% and 15.0%. Egg
parasitism is not known to have exceeded 27.0%. The associated factors
are the relatively low density of S. litura on this experimental farm (indi-
cating that only the parasites with a highly efficient searching ability will
have any impact) and the effectiveness of the insecticide applications.
Birds are known to predate S. litura larvae on the ICRISA T research
farm. Cattle egrets removed 62% of the larvae released into un netted
enclosures (compared with netted enclosures). This may be an exagge-
ration of what happens in farmers' fields because the birds have learned
that they may get food where they see the enclosures on our research farm.
Cattle egrets are certainly present in farmers' fields in Asia and Australia
and are known to include noctuid caterpillars in their diet (Siegfried,
1971). There is a need to learn more about the role of these and other birds
as predators in groundnut and other crops.
S. litura is susceptible to viral diseases. We have heard verbal accounts of
farmers in India spraying diluted suspensions made from diseased larvae to
their crops. However, we have no details of the larval equivalents per unit
area or the effectiveness of such activities.
Shanower et al. (1992) list 38 species of parasites known to be associated
with the groundnut leaf miner. Seventeen of these were reared from larvae
collected on the ICRISAT farm, and approximately half were known to be
primary parasites. There were changes in dominance among parasite
species during a season. For instance, in the post-rainy season 1987-1988,
Sympiesis dolichogaster emerged from 26% of the parasitized larvae of the
first groundnut leaf miner generation and declined to 12% and 16% by the
third and fourth generation. Stenomesius japonicus emergences increased
from 6% to 22% during the same period. The other species involved,
which also emerged at higher rates as the season progressed, were
Goniozus sp., Chelonus sp. and a group of three braconids - Apanteles,
A vga choaspes and Bracon sp. The 'other species' group, at 40%, predomi-
nated by generation 4 and included unidentifiable species, some of which
could have been hyperparasites.
This study also showed that:
• parasitism levels were not influenced by the genotypes - ICG 1697
(NCAc 17090) and ICG 799 (Kadiri 3 = Robut 33-1);
• insecticide application (dimethoate at 240 g/ha in 350 I water, once early
in each caterpillar generation) reduced parasitism levels by 7-14%;
and
• diseases accounted for up to 30% of the larval mortality (a new finding).
The latter observation indicated that a degree of larval mortality remained
unaccounted for. This was assumed to be predation by carabids, e.g. Chlaenius
sp. (Shanower and Ranga Rao, 1990) and spiders. The remaining
448 Groundnut pests
TABLE 11.18 Influence of groundnut genotype on the
rate of groundnut leaf miner parasitism at ICRISA T
Center, rainy season 1992 (d = host density, % =
percent parasitism) (data collected by Research Fellow
M. L.J. Sison)
Genotype Leaf miner Parasitization I

larvae per m 2 (%)
(d)
ICG 221 42 16
ICG 156 28 26
ICG 5240 26 26
ICG 2271 21 36
ICG 5040 78 80
ICGY 86031 31 22
ICG 799 51 30
ICG 5044 154 26
SEm ± 13.1
I Sample size: 50 larvae in each treatment.
questions about the interactions between host density, primary and sec-
ondary parasites and the mortality caused to all three levels by a range of
insecticides, possibly applied at different times during the life-cycle of the
caterpillar, remains as an example of a research project revealing further
sets of problems. The contribution of the hyperparasites to the population
dynamics of the herbivore is an important issue because we believe that the
maintenance of a high parasitism rate (which can exceed 90%) is a lead
factor in the management of the groundnut leaf miner.
Although Shanower et al. (1992) found that host plant genotype had no
effect on the rate of parasitism (all species), this is not necessarily always
the case. Observations at ICRISA T Center, on crops growing in an area
that has never been treated with insecticides, indicate that host genotype
and species can influence the level of groundnut leaf miner parasitism.
Table 11.18 shows that, in a comparison of four genotypes, the rate of
groundnut leaf miner parasitism was particularly low on aphid and jassid
resistant ICG 5240, which is not known to be resistant to this caterpillar.
Only ICG 156 is recognized as having a degree of resistance to the
groundnut leaf miner.
Turlings and Tumlinson (1991) provide information to indicate that it is
possible for chemicals released from a plant as a result of herbivore
damage to attract parasites to the scene of the activity and thus to their
host(s). There is a good case for following up this matter for groundnut in
view of the clear effect of genotype on ground nut leaf miner parasitism.
The rate of parasitism (all species) also proved to be higher by 8% or
TABLE 11.19 Density of groundnut leaf miner larvae and larval parasitism on sale
(s-) and intercropped (i-) groundnut (gnt) and soybean (soy/: fCRfSAT Center,
data for 22 August (rainy season)
s-gnt s-soy i-gnt i-soy ±SE

Larvae per 25 plants 3.1 4.7 3.3 6.3 1.3
Larvae/m 2 soil
surface 46.0 183.0 52.0 243.0 22.0
Larvae/lOO cm 2 leaf 9.1 19.4 9.7 26.7 1.4
% parasitism 36.0 47.7 31.4 38.7 2.6
'Plot size = 15 x 15 m, with seed sown in four rows on 1 m wide raised beds. Alternate beds
of the two crop species were sown in the intercropped treatments. There were five replicates
of the three treatments.
12% on soybean than on groundnut, depending upon whether the crops

were sole crops or intercrops (Table 11.19). Host density was higher on the
soybean, so that the higher rate of parasitism could be a function of the
parasites' searching efficiency, i.e. under the relatively low host density in
this experiment, the parasites were able to sting more hosts on soybean
because they needed to search fewer leaves to find a caterpillar - a complex
aspect of parasite ecology in its own right (Hassell, 1982).
11.5.3 Africa
Weaving (1980) found that up to 56% of the eggs of Hilda patruelis are
parasitized by Psyllechthrus oophagus (Hymenoptera: Encyrtidae).
However, the rate drops considerably during the rainy season, which is
when groundnut is grown.
Colonies of Aphis craccivora living on newly established groundnut
crops (1--4 weeks after emergence) in Malawi were virtually without excep-
tion accompanied by (unidentified) syrphids, coccinellids and lacewings.
Aphidioid mummies were also present. The aphid colonies died out within
a week of this observation. The impression, as no data were collected, was
that the predators were mainly responsible for this decline in aphid popu-
lation density (observation by Wightman).
Later in the season, Wightman also noted the presence of high densities
(up to one per 10 plants) of mantids in groundnut crops throughout
southern Africa. They were of sufficient size «7 em body length) and
density to warrant an investigation of their contribution to the natural
control of potential groundnut pests in Africa.
The predominant arthropod life form, however, appeared to be ants.
Members of genera such as Pachycondyla, Myrmiccaria and Platythyrea
were often abundant in groundnut fields and were seen in Malawi and
Tanzania carrying caterpillars and other prey to their nests. Reimer (1988)
450 Groundnut pests
found that thrips were eaten by ants (and anthocorid bugs, which are also
common in groundnut fields in southern Africa).
Even less is known about the natural control of soil insects. In Malawi,
ants were seen dragging a dropped live \vhite grub larva, weighing approxi-
mately 3 g, along the furrow between two rows of groundnut plants. It is
not known what happens when such an en:counter between ant and white
grub takes place underground. Wireworms are also recognized as the
natural enemies of white grubs, but they can damage pods as well.
Ants are the natural foe of termites (Logan et at.. 1990). Wightman and
Wightman (1988) found that the admixture of insecticides with the soil in
groundnut fields can disrupt ant activity, whilst leaving termites unaffec-
ted. Whilst sampling the soil of groundnut fields for insects, Wightman
found larvae belonging to the dipteran families (unidentifiable to species
from larvae by the British Museum of Natural History) Scenopenidae,
Mydidae and Therevidae. They are all known to prey on either Coleoptera
larvae, per se, or arthropods and earthworms (Therevidae), and may,
therefore, include white grubs in their prey.
A fungal disease, Metarhyzium anisopliae, the green muscardine fungus,
is currently under evaluation for the control of the white grubs that attack
groundnut in Queensland (Milner, Rogers and Brier, personal communi-
cation; Milner, 1989, 1992). After many years of evaluation, the selection
of strains that are highly pathogenic to scarab larvae has allowed the
application of this technique to proceed to the commercialization phase for
pasture and sugar pests. The propagation and dispersal of the fungus is
relatively simple. The successful outcome of the trials in Australia and the
extension of the technology to Asia and Africa is the kind of breakthrough
needed to give leverage on the almost intractable white grub problem.
11.5.4 Comments on the natural control of insects living on groundnut

crops
It is known that the potential for natural control to make a major contri-
bution in maintaining the herbivores living on groundnut at levels lower
than action thresholds is high. Unfortunately, considerably less is known
about the dynamics of the various processes. This is especially true of
predation, where there is ample scope for applying immunological tech-
niques to determine the nature and quantity of the prey of suspected
arthropod predators (e.g. Giller, 1984; Stuart and Greenstone, 1990).
Presumptive vertebrate predators could be studied by more conventional
techniques involving trapping and faecal analyses.
11.6 INSECTICIDES
We are, in general, somewhat diffident about emphasising the application

of insecticides for reducing the density of foliage feeding insects, except at
Insecticides 451
the seedling stage, and perhaps to the soil. The data available indicate that
mirids, as destroyers of flower buds, are also an exception and, where
feasible, should be eliminated if present in crops during the flowering
stage, at least in the more determinate varieties.
Since 'peanut entomological literature is replete with the effectiveness of
insecticides in reducing pest populations and to increasing peanut yield,'
(to quote Lynch and Douce, 1992), we shall not go into the details of the
many insect/insecticide options. We shall instead look at areas that are not
covered so well.
Although no documentation referring specifically to groundnut has been
located, the 'Green' environmental lobby in the USA is putting pressure
on the agroindustry as a whole to reduce pesticide usage. Without entering
the rights and wrongs of such matters, this will undoubtedly push the
groundnut industry towards exploiting the resistance that already exists in
NC 6 and the genotypes and species discussed in section 11.4. It is
fortuitous that excellent co-operation between discipline scientists in the
south-east of the USA has led to the identification of such a choice of
material. This review also signals an increase in interest in the process of
natural control among groundnut entomologists in the USA since the late
1980s.
Our impression is that A. hypogaea is naturally resistant to many insects
and that many of the insect pest problems on groundnut are induced by
insecticides (or suboptimal management procedures). Tait and N apompeth
(1987) provide many examples of how insecticide use in the less developed
countries has created more problems than it has solved. Circumstantial
evidence for this is Wightman's observation, during a survey of groundnut
fields in southern Africa. that he only found significant defoliation (by
Helicoverpa sp. and Spodoptera littoralis) on two research stations where
insecticides had been applied in response to minor defoliation.
Amin (1988) stressed that, up to the mid 1960s, there were many fewer
insects recognized as groundnut pests in India than there are today. This
must be attributed to changes in groundnut management, and one such
change is the increase in insecticide application.
Ranga Rao and Shanower (1988) found that 70% of farmers visited in a
post-rainy season survey of groundnut fields in Andhra Pradesh, India, had
applied insecticides to their ground nut crops, mainly in response to the
appearance of groundnut leaf miner. Heavy users had crops with bad
defoliation by S. litura and Helicoverpa armigera, but their crops were in
no better condition than neighbours who had not applied insecticides. The
latter had little defoliator injury but may have had reduced yields (though
not necessarily submaximal profits) as a result of groundnut leaf miner
activity. This was attributed to the destruction of the natural control
process by insecticides. Our experience at ICRISAT is that areas of
groundnut fields that have had insecticides applied to them always suffer
more from defoliators (Table 11.20). The failure by farmers to eliminate
452 Groundnut pests
TABLE 11.20 Damage by defoliators to groundnut in plots treated and untreated
with insecticides: consolidated data from fCRfSAT Center (experimental plots) in
post-rainy season only
Percentage of leaflets with defoliator damage
With insecticides No insecticide
1984-85 7.0 (8 x dimethoate) 0.3

1985-86 9.0 (8 x dimethoate) 5.3
1986-87 20.7 (8 x dimethoate) 5.8
1987-88 5.7 (2 x monocrotophos) 2.9
1990--91 1.9 (2 x monocrotophos) 0.3
the target Lepidoptera implies that these defoliators were no longer

susceptible to the insecticides (carbamates, organophosphates and
pyrethroids applied singly or as ad hoc cocktails). Insecticide resistance in
both of these species in this area has been documented (Ramakrishnan
etal., 1984; Armesetal., 1992).
Unfortunately, it is not possible to rerun history and perform controlled
experiments on such matters, but it is possible to attempt to reverse the
process. Prior to 1984, insecticides were applied to groundnut crops on the
ICRISAT farm in response to the perceived needs of the scientists
(breeders, physiologists etc.) involved, reacting to what was often minor
damage. This resulted in a treadmill effect, to the extent that eight or nine
applications were being made per season (D.S. Bisht, personal communi-
cation). In the 1984 post-rainy season this process ceased and insecticides
have only been applied at the behest of a groundnut entomologist, accord-
ing to thresholds that have been slowly relaxed. The result is that there has
been a steady decline in the average number of applications made per
season and the number of pest outbreaks (Table 11.21). The unfortunate
result is that it is now difficult for breeders, entomologists and virologists to
screen for resistance in realistic conditions. Table 11.21 indicates a reversal
in the downward trend in the amount of insecticide applied to ground nut in
the 1989 rainy season. This corresponded to increased levels of insecticide
application in response to heavy outbreaks of H. armigera on chickpea and
pigeon-pea that probably diminished the level of the natural control
process across the whole farm.
11.6.1 Flower pests

What is not shown in Table 11.21 is that, often, only one low rate
application of dimethoate was applied at the start of a season (5-12
Insecticides 453
TABLE 11.21 Insecticide (kg or I formulated product) applied to groundnut fields
on the ICRISAT Center Research farm (source D.S. Bisht, Farm Manager)
Season Total Total Mean Materials

area applied ha- I
(ha) (kg or I)
Up to 20-30 >200 c.9

1984
1984 Rainy 20.00 102.10 5.10 Di, En, Dv, Me
Post-rainy 30.00 123.50 4.12 Di, En, Ca, Dv
1985 Rainy 13.90 50.00 3.57 Di, En, Dv, Me
Post-rainy 29.73 65.75 2.19 Di, En, Dv, Me
1986 Rainy 22.00 44.63 2.03 Di, En, Dv, Fe, Lo, Me
Post-rainy 31.41 31.20 1.00 Di, En, Me, Fe
1987 Rainy 29.90 32.00 1.07 Di, Dv
Post-rainy 34.75 36.25 1.00 Di,Dv
1988 Rainy 17.14 20.57 1.20 Di
Post-rainy 26.90 31.55 1.17 Di
1989 Rainy 20.00 66.80 3.30 Di, Qu, En, Ca, Dv
Post-rainy 19.95 36.5 1.92 Di, Qu, Dv
1990 Rainy No data
Post-rainy 24.00 62.4 2.60 Di, En
Ca = carbaryl, Di = dimcthoate, Dv = dichlorvos, En = cndosulphan, Fe = fenvalerate,

Lo = Lorsban, Me = metasystox, Qu = quinalphos
days after emergence) to kill thrips and jassids. Whilst even this may not
have been necessary in economic terms, it served certain cosmetic needs of
a research station. It is suspected that this regime would eliminate the
mirids, which attack early and can have a marked impact on the flowering,
without jeopardizing the natural control process. This is because the major
potential defoliators have not established themselves by this stage so that
there is no attendant cohort of parasites and predators. (These comments
arise from discussion with D.l. Rogers and H.B. Brier, Queensland
Department of Primary Industries.)
11.6.2 Soil insects

Currently there is no proven alternative to insecticides for the control of
soil insects, other than the promise presented by certain resistant geno-
types and wild species (NC 6 against Diabrotica is the exception).
Elasmopalpus remains a problem insect in the USA, especially in
dry seasons, but can be controlled by granular formulations of a range
454 Groundnut pests
of insecticides (Smith and Barfield, 1982). However, as they are best
incorporated in the soil prior to sowing, they must be applied prophyl-
actically. This is also the conclusion arrived in India for the control of
white grubs where ph orate is the preferred insecticide (Yadav, 1991 publi-
cation), although Bakhetia (1982) found that seed dressings of other
insecticides were effective for white grub control. The best yield response
was achieved when the insecticide was mixed as seed dressing with thiram,
for the control of Aspergillus niger, the cause of collar rot (Bakhetia et al.,
1982).
Logan et al. (J992) sought alternatives to cyclodiene insecticides for soil
insect control by comparing the effects of aldrin, chlorpyrifos and carbosul-
phan on termites, white grubs and other insects living on or under ground-
nut stands in India and the Sudan. Chlorpyrifos was, in general, the most
effective in that it gave as good results as aldrin, especially in a slow release
formulation. The only problem was that residues were found in the seeds
after harvest. Isofenphos granules were not as effective as chlorpyrifos for
controlling soil insects, but could have a place in pest management pro-
grammes because the systemic effect of this material considerably reduced
the incidence of groundnut leaf miner caterpillars.
11.7 CULTURAL CONTROL
The concept of cultural control covers all management activities a farmer

could execute that are not included under the headings of the preceding
three sections. At the same time, cultural control may also include these
activities because they are very much intertwined and invariably involve
management decisions. It is when we come to this area that we should
begin to be able to consider catalysis in the form of insect densities
stabilized below economic levels resulting from the simultaneous appli-
cation of two or more pest management tools. However, before integration
is considered it is necessary to review some of the management options that
could be considered by groundnut farmers.
11.7.1 Intercropping and habitat diversification

Risch et al. (1983) reviewed the relationships between agricultural diversity
and pest incidence. They concluded that the benefits, in terms of species
abundance and outbreak suppression, were more likely to be associated
with interference with insect dispersal activity than enhanced natural con-
trol. This may point to the need to design systems specifically to assist the
proliferation of natural enemies. This means providing food (nectar
sources) for parasitoids and refuges for arthropod predators (sometimes
called 'weeds'), and considering an agroforestry component in the farming
Cultural control 455
system to encourage birds, apart from the other benefits. This is why we
stress that there is a close link between cultural control and natural control.
Low (or no) technology farmers have diversified their farm systems
probably since farming began, largely to ensure that they have something
to harvest at the end of the season. Sowing groundnuts between rows of a
cereal (maize, pearl millet or sorghum) is common in parts of Africa but is
not apparently practised so often in Asia. The indications are that the
density of thrips, jassids and groundnut leaf miners on groundnut is
reduced by this practice (Wightman and Amin, 1988; Muthiah et al., 1991).
Farrell (1976c) found that, in Malawi, a groundnut-bean (Phaseolus)
intercrop reduced the incidence of GRV because the aphids became
hooked on leaf hairs on the bean leaves as they moved from one row to
another. Farmers in southern India sow castor at a low density in ground-
nut fields to attract female Spodoptera moths to its leaves for oviposition.
The egg masses are easily detected on castor leaves and can be destroyed
by hand.
It is our experience that cowpea and groundnut are not good com-
panions because of the risk of the spread of Aphis craccivora from cowpea
to groundnut. Also, the practice of juxtaposing soybean and groundnut,
which is common in Indonesia, could lead to a bad attack of groundnut leaf
miner in the groundnut crop. This insect prefers soybean (Table 11.19) but
can transfer if the time of soybean harvest corresponds to the pupal or
adult stage. We exploit this process by growing 1 m wide beds of soybean
around our experimental plots. They are pulled up and left to dry on the
soil surface during the pupal stage so that the moths leave the dying plants
and oviposit on the groundnut plants.
A stand can also be diversified by sowing more than one genotype of the
same species in alternate rows or beds, as a mixture or in some other
pattern. To our knowledge, this process has not yet been attempted with
groundnut, but it appears to be viable in chickpea where a high yielding but
pest susceptible variety was 'protected' by a lower yielding variety with
high pest resistance (ICRISAT, 1992).
11.7.2 Mulching
(a) Organic mulches to protect the harvested product

Farmers in Africa commonly windrow the newly harvested plants to permit
sun-drying. Odontotermes frequently penetrate the piles of drying plants
and damage the haulm and seeds. This can reduce the pod yield by as much
as 30% or 40% and, of equal importance, increase markedly the sub-
sequent levels of aflatoxin contamination.
Jpomaea fistulosa (morning glory bush) is a common roadside plant in
India. Unlike most plants in this environment, the leaves are not eaten by
insects or goats. Observation of this fact led to a possible approach
456 Groundnut pests
TABLE 11.22 Effect of organic mulches on the percentage
and level of pod scarification by termites (Odontotermes sp.
and Microtermes obcsi)
Pods Scarification
scarified mean score
(%) (0-4)
Neem cake 2 (0.02) 0.04

lpomaea 7 (0.07) 0.14
Celosia 17 (0.17) 0.38
Sunn hemp 59 (0.66) 1.69
Bare ground control 36 (0.37) 0.88
SE ± (0.02) 0.06
F-value 156.4** 118.7**
Plot size = 200 x 50 em

20 replicates
Neem cake mulch = 2.5 cm thick; others = 5 em thick
Scoring: 0 = no scarification
1 = 1-25% of shell surface scari,ficd
2 = 26-50%
3 = 51-75%
4 = 76-100%
(Figures in parentheses are arcsine-transformed value of radians)
to protecting pods as they dry on the ground after harvest. The idea of
using a mulch made of the chopped Jpomaea leaves and stems was ex-
tended to the possibility of using other plants that are not attacked by
termites for the same purpose. The results (Gold et al., 1989; Table 11.22)
indicated that the Jpomaea mulch and neem cake gave drying pods a high
level of protection from termites. A mulch made of Celosia argentea, a
common weed that survives in the vicinity of areas of high termite activity
on the ICRISAT farm, gave some protection, but sunn hemp increased the
level of termite activity, compared with bare ground. A subsequent experi-
ment (ICRISAT, 1991a) indicated that the long-term benefits of using
Jpomaea to discourage the surface activity of termites were negligible
compared with a neem cake mulch. A neem cake mulch applied to row
crops was then shown to have no beneficial effect on yield or foliar insect
management - and was very costly (Ranga Rao et al., 1991).
The data available indicates that selected organic mulches could reduce
the termite and aflatoxin problem associated with field drying that has
concerned farmers and traders alike for many years. There is no evidence
to link such mulches with benefits to the growing crop. However, our
experiments were not exhaustive in this respect and there is ample scope to
investigate the link between enhanced crop water economy (reduced soil
evaporation) and weed management associated with mulching and the
ability of the plant to tolerate insect attack.
Grainge et al. (1985) indicate that Jpomaea spp. have anti-fungal and
anti-insect (bruchids, 'bugs', various Homoptera and flea beetles) proper-
ties. This is clearly a fertile area for further investigation.
(b) Synthetic mulch to reduce virus vector incidence

The ability of a shiny (reflective) or white mulch to protect crops from
aphids and the viruses they transmit has been known for many years. The
theory and practice are well established (Kennedy et al., 1961; Kring, 1972;
Smith and Webb, 1969). Winged aphids respond positively to a short-wave
radiation source (such as the sky) when they are ready to initiate their
dispersal flight and whilst they have sufficient stored energy to sustain this
flight. When 'fatigued' or 'hungry' they react positively to long-wave
radiation such as that reflected by vegetation. Thus, whilst they are in the
early phases of their flight they respond to the major source of short-wave
radiation - the sky - but are confused and repelled when they receive a
(reflected) sky signal from the ground when in the end-of-flight host-
seeking stage. Pr.esumably, they then fly on to the field of a non-mulching
neighbour.
When it comes to protecting a crop from a non-persistent virus (e.g.
peanut stripe virus) there are only two options: growing a resistant variety,
if one is available, or preventing the vector coming into contact with the
crop. Clearly the information about aphids' flight physiology is of funda-
mental value for protecting a crop from a non-persistent virus, which can
be transmitted during just a few seconds' probing by a viruliferous aphid.
With persistent viruses (e.g groundnut rosette virus) there are more op-
tions but avoidance is still one of them (A'Brook, 1964).
Xu Zeyong et al. (personal communication) have shown that plastic
mulches may be a viable method of protecting groundnut crops from the
non-persistent peanut stripe virus, for which farmers currently have no
satisfactory management procedure. The data so far available (Table
11.23) indicate the potential benefits of this approach in terms of reducing
the number of aphids alighting in a crop, virus incidence and yield, even
though the reporting years were quite different climatically. Dr Xu Zeyong
has indicated that the plastic material is available at prices that are within
the economic bounds of groundnut farmers in the People's Republic of
China. The benefits in yield increase indicated in Table 11.23 are almost
certainly related to the lower virus incidence but may also be associated
with lower levels of fungal disease (some foliar diseases are initiated by
spores splashing up from the soil surface onto the leaves), weed manage-
ment and a reduction in the amount of water evaporating from the soil
surface. These matters are open for further evaluation.
Reflective mulches or aluminium strips painted on a black plastic mulch
have recently been shown to protect narrow-leafed lupin and summer
squash from non-persistant viruses by repelling the aphid vectors (Jones,
458 Groundnut pests
TABLE 11.23 Influence of sheets of silver and transparent plastic used as a
mulches on the number of aphids caught in yellow pan traps placed in the plots,
peanut stripe virus (PStV) incidence and yields in field trials in Wuchang, Peoples
Republic of China, communicated by Dr Xu Zeyong, Oil Crops Research Institute
Silver plastic Transparent Bare soil

mulch plastic mulch control
Yellow pan catches

(aphids per season)
1990 37 31 268
1991 5 59 404
Virus incidence (%)
1990
8 weeks after sowing 14.7 18.1 37.7
1991
Yield (t/ha)
1990 3.5 3.6 2.7
1991 1.8 2.1 0.7
1991; Lamont 1990). Shiny plastic mulches are used in Thailand (for
instance, in vegetable seed nurseries) for weed control and water
conservation.
It is likely that cultural practices will need to be modified to accommo-
date a reflective mulch if they include recommendations to ensure rapid
canopy closure. This is because the mulch can only work if a sufficient area
is exposed to the sky. However, as we anticipate that the mulch will also
act as a weed control agent, this should not matter. Early canopy closure is
usually recommended to reduce the number of weedings needed - and to
make the crop less attractive to immigrant aphids, a role we anticipate will
be taken on by the shiny mulch (A'Brook, 1964; Farrell, 1976a).
This is a good example of why and how pest management should be
approached in a multidisciplinary manner. The preimplementation
(research) phase needed for the further development of the shiny mulch
technique clearly needs input from agronomists, weed scientists, virologists
and entomologists.
11.7.3 Irrigation management

The likes of Smith and Johnson (1989) make it clear that the lesser corn
stalk borer is not likely to achieve pest status under irrigated conditions.
The same is partially true of the groundnut leaf miner (Wheatley et al.,
1989) but not for the groundnut aphid, which proliferated in the wetter end
of the drought stress gradient despite being pounded by water from an
overhead irrigation system (Ranga Rao et al., 1991c).
It also appears that the source of irrigation is important. The only
outbreak of red spider mite on the ICRISA T farm in recent years was
associated with furrow irrigation combined with insecticide application.
Neighbouring fields, which were either irrigated with sprinkler and sprayed
with a similar insecticide regime or irrigated with a sprinkler and received
no insecticide, had no mite outbreaks (densities ranged from 13-110 mites
per 50 leaflets). Overhead irrigation reduced the mite density from 6323 to
1282 per 50 leaflets in the outbreak field in a matter of days (Ranga Rao et
al., 1990) without recourse to pesticide application. This reduction was
sufficient to allow the labourers to harvest without suffering from skin
irritation caused by the mite.
11.7.4 Avoidance
Groundnut, being an annual or short season crop, depending upon the
genotype and where it is grown, relies mainly upon immigration to provide
its quota of foliage feeders. (Possible exceptions are the noctuids and hairy
caterpillars that aestivate or hibernate as pupae in the soil.) In line with our
data indicating that the older a crop is, the less likely it is to be damaged by
insects, it would appear to be a rule of thumb to sow groundnut crops
early, before pests have a chance to multiply on their winter/hot/dry season
hosts and start the migration process. This is by now conventional wisdom
for groundnut in southern Africa, where crops sown with the break in the
dry season avoid groundnut rosette virus (section 11.7.2(b)). It is likely
that co-ordinated early sowing by government behest in Malawi is respon-
sible for the virtual elimination of this disease in that country.
A similar phenomenon exists in the USA where the corn earworm passes
from corn to groundnut, soybean and cotton in late July and August.
Sowing the crops early (early to mid April) diminishes the damage caused
by this insect (Lynch and Douce, 1992). Early sowing is also likely to
reduce the risk of damage being caused by the lesser corn stalk borer
(Mack and Backman, 1990).
Our experience with thrips at ICRISAT is mixed. It is not unusual for
early sown crops to be badly affected by bud necrosis virus and later sown
crops to be unscathed. At the other end of the season, early or timely
harvest is called for to avoid termite attack and to reduce the time that the
mature crop is exposed to other pod-eating pests, including mammals
(Lynch et al., 1986). A delay in harvest is therefore likely to increase
aflatoxin incidence.
Some cultural practices are believed to promote pest outbreaks. For
460 Groundnut pests
instance, the development of irrigation systems in many areas of the semi-
arid tropics during the last 30 years has allowed farmers to extend their
operations beyond the rainy season. This unquestionably has had a major
positive impact on agricultural production. Unfortunately there are some
negative effects that would not have been anticipated by the planners. For
instance, Yadav (1981) notes the proliferation of white grub damage in
northern India where supplementary irrigation prevents soil drying out.
Desiccation and soil heating as a result of drought probably affect the grubs
more than groundnut plants.
Until perhaps 30 years ago, in southern India Spodoptera litura was
limited to being a pest of tobacco. It is presumed (Amin, 1988) that its
adaptation to groundnut and other crops in southern India was a result of
its being exposed to these erstwhile single season crops for more of the year
than was the norm under rainfed conditions. It is not unusual to see crops
at all stages of development from June to March. A short close season
(April to May) gives some respite, except in parts of Tamil Nadu where
groundnut is grown all through the year. Groundnut leaf miner and
Spodoptera litura are particular problems in these areas, as is Helicoverpa
armigera, which has only recently been recognized as another (induced)
pest of groundnut in southern India.
11.7.5 Soil preparation and related cultural practices

Yadav (1981) indicated that deep ploughing can reduce white grub
damage, especially during the pupal stage. The 'mode of action' clearly
involves the physical damage to the insects as well as disruption of the
grubs' environment and their exposure to the sun. Birds are often seen
following the plough, presumably to pick up exposed insects (Syamsunder
Rao, 1992). The practice of ploughing in the cool of the night, which is
made possible with the introduction of the tractors that are a characteristic
of the 'white grub belt' of northern India, does not contribute to white grub
control. Firstly, birds do not hunt at night; secondly, the grubs are not
exposed to the burning and desiccating effects of solar radiation.
Mack and Backman (1990) compared the effects of conventional tillage
(ploughing and disking before sowing), reduced tillage (sowing into wheat
stubble with a combined subsoiler and seed drill) and 'no-till' (sowing into
burned wheat stubble) on insect densities. The tillage system had no effect
on the population density of the pest and beneficial insects sampled.
Farmers in Nigeria said they were able to control termites by a number
of processes: drumming; burying dead animals, cassava meal, fish guts
or the contents of torch batteries in their fields; and introducing soldier
ants into the termite nests (Malaka, 1972). There were several other
methods involving growing plants or their extracts (Sanseveria libericum,
Ocimum basilicum, Parkia clappertoniana, Vetivera nigritana, Digitaria
Research related to insect pests of stored ground nut 461
sp., Cymbopogon sp., and Pennisetum purpureum) that are worth evalua-
tion.
11.7.6 Other methods

Farmers may use a wide variety of other methods of managing their pests
that may be beyond the ability of scientists to evaluate under controlled
conditions. Such practices include making smoky bonfires around fields to
disrupt the evening post-eclosion flights of Amsacta spp., burning hand
picked blister beetles on fires lit close to fields (the beetles may release
an alarm pheromone) and collecting and killing cockchafers during the
crepuscular feeding and mating forays to specific trees. Such practices,
when combined with other cultural activities, may help to sustain insect
densities at levels below which they become pests. The benefits of other,
technology based approaches, such as trapping noctuid moths in ultraviolet
light traps, releasing egg parasites or catching male moths in pheromone
traps, have also yet to be proved of value.
A new approach to aphid control has been described by Harrewijn et al.
(1991). They achieved perfect or almost perfect protection from insects
and the vectored virus diseases of potato crops by covering the growing
plants with an ultrafine polypropylene net. The management of non-
persistent viruses, such as the peanut stripe virus in Asia, remains an
unsolved problem, especially, as in this case, the virus is seed-transmitted.
Clearly, the technique of covering crops with such a fine web, which is still
under investigation in Holland, could become a viable protection method
for farmers' groundnut crops in Asia. However, details of cost and avail-
ability have not reached us and we suspect that both could be constraints to
its widespread adoption. The technique may have a special application in
seed or research farms because of the need to provide virus-free seed for
farmers and researchers.
11.8 RESEARCH RELATED TO INSECT PESTS OF STORED

GROUND NUT
A number of insects feed on stored groundnut but perhaps only one is

specifically associated with this product, at least in Africa and Asia -
Caryedon serratus, the groundnut bruchid, borer or 'weevil'. The biology
and management of this and the other species have been discussed in detail
by Dick (1987b) and Wightman et al., 1990. This section simply updates
what has gone before.
Ranga Rao et al. (1987) reported that a black carpet beetle, Attagenus
fasciatus (Dermestidae), was found in groundnut stores in India. This
species is one of the few capable of penetrating the pod (as a neonate).
462 Groundnut pests
Most groundnut storage pests gain access to the seed of the unhulled
product via broken shells and testa.
11.8.1 Resistance
One way of limiting the activity of such storage pests is to seek genotypes
with resistance factors in the shell. Entomologists have located such geno-
types and it is for others to decide whether to attempt to breed such traits
into commercial varieties.
The pyralids Plodia interpunctella (the Indian meal moth) and Cadra
cautella (the almond moth) are of particular concern in commercial and
farm stores in the south-eastern states of the USA. Kashyap and Campbell
(1990) tested 39 A. hypogaea hybrids (including established varieties) for
resistance to these species. They found, with a little variation, that breed-
ing line 1O-PlO-BI-BI-BI-BI-B2, variety NC 7 and entries with NC 343,
NC 2214 and NC 2232 in their parentage were among the most resistant to
both species. Assays were made of oviposition preference, larval develop-
ment and damage.
Advanced breeding lines have been screened for resistance to Caryedon
serratus (256 lines) and Corcyra cephalonica (306 lines) at ICRISAT
Center to discover whether a given line is likely to be more or less resistant
to storage pests (Mittal, Wightman and Dwivedi, unpublished). Most
varieties tested were neither particularly resistant nor susceptible to either
insect. Seventeen were more resistant than the most resistant control to
Caryedon serratus and 26 had resistance to Corcyra cephalonica.
11.8.2 Protecting stored seed

Farmers in developing countries often have to pay a large premium when
buying groundnut seed because they are not able to store their own seed
from one season to the next without loss of quality as a result of mould and
insect infestation. An attapulgite-based clay dust applied to groundnut
pods (0.5% w/w) limited the ability of Corcyra cephalonica to reproduce to
any significant extent (Mittal and Wightman, 1989). This was not a total
surprise as the principle that an inert dust can protect stored food stuffs
from insects has been demonstrated for a number of scenarios (Shawir et
al., 1988). As fungi also reduce the quality of stored seed, we substituted a
dust formulation of a number of fungicides (at 3 g/kg) for the clay dust.
C. cephalonica did not propagate with these treatments (ICRISAT, 1991,
pp. 87-88). Caryedon serratus was not as responsive to the fungicide dusts,
in general, as it was to the attapulgite clay dust. However, the formulation
of thiram (75 WDP) that was tested could be substituted for the dust. This
experimentation was not continued in on-farm conditions but the infor-
mation was made available to the Indian National Agricultural Research
Program and to the general public via a newspaper article stressing that
Integrated control of groundnut pests 463
seed from pods treated with a fungicide should not be eaten unless
thoroughly cleaned.
11.9 INTEGRATED CONTROL OF GROUNDNUT PESTS
We define integrated pest management as one or more activities that are

carried out by farmers that result in the density of potential pest popu-
lations being maintained below levels at which they become pests, without
endangering the productivity of the farming system as a whole, the health
of the farm family and its livestock, and the quality of the adjacent and
downstream environments.
One such farmer activity could be sowing an alternative to a particular
crop because the latter could not be harvested without using excessive
amounts of pesticide or without a high risk of suffering excessive crop loss.
Examples in the case of groundnut are few but include the situation in parts
of Tanzania and Malawi where this crop is not grown because of the risk of
hilda attack.
The keys to the management of insect pests of groundnut are minimizing
insecticide application (or, in the case of many African farms, maintaining
the current zero to minimal level) and exploiting host plant resistance,
combined with cultural practices, especially those that enhance natural
control processes. For instance, Campbell and Wynne (1985) demon-
strated that NC 6, which has resistance to thrips, jassids, Heliothis zea and
Diabrotica undecimpunctata howardi, can be grown without yield loss
penalty (in North Carolina) with no or minimal insecticide application -
compared with Florigiant.
Unfortunately, there may always be pests that will cause damage in
certain years. Examples are Elasmopalpus lignosellus in the USA and the
groundnut leaf miner in Asia. Perhaps it is only coincidence but the
biological success of both of these insects is favoured by hot, dry con-
ditions. Both can be controlled with granular insecticides incorporated into
the soil at sowing, i.e. well before a pest attack can be forecast. The answer
to the management of these insects may have to wait until farmers can rely
on long-term (6-8 week) weather forecasts.
11.9.1 IPM for groundnut farmers - current and future prospects

As has been made clear, we do not exclude insecticides from rPM. If this is
generally accepted, it follows that pest management specialists are obliged
to provide farmers or their advisors with the means of deciding when to
apply insecticides selectively and which are most appropriate to alleviate or
avoid a given condition. Such a scheme has been in place in Georgia
(USA) for over a decade (Douce, 1982). It involves information exchange
at farmer meetings, publications, field scouting and pheromone traps for
464 Groundnut pests
monitoring key insects. Lynch and Douce (1992) indicated that, in its early
stages, participants benefitted by a yield bonus of nearly 1 t/ha (27%) for
an increased outlay of $22/ha per season for pesticides. A more recent
evaluation showed that the major participants were spending less on
pesticides than non-users and were maintaining high yields.
This indicates two features that should be components of well run,
extension lead IPM schemes: routine monitoring of impact of recommen-
dations, and continuous attempts to improve the advice provided, espe-
cially if faced with a labile pest scene.
The development of IPM schemes for groundnut is not limited to
developed countries. In the Philippines (San Mariano, Isabella) an IPM
programme involving pesticides, an egg parasite and Bacillus thuringiensis
and resistant variety BPI Pn-9 gave higher yields with less pesticide appli-
cation than no treatment controls and farmers' practices (Campbell et al.,
unpublished).
The process of developing a similar programme for southern India has
been initiated. In many districts there is a clear seasonal sequence of
events:
1. Farmers apply insecticides early in the season in response to jassid and
thrips symptoms.
2. This reduces the density of parasites and predators (including birds
that will only stay in or around a farm if there are at least a few insects
to eat).
3. Because of 2, groundnut leaf miner densities increase.
4. Farmers apply insecticide to kill the leaf miners.
5. The level of natural control is reduced even further.
6. Outbreaks of Spodoptera lilura appear.
7. More insecticides are applied.
8. At this stage insecticide resistance almost certainly occurs so that
higher concentrations, cocktails or over-frequent applications have
been reported.
9. More severe S. litura outbreaks occur, accompanied by the appearance
of Helicoverpa armigera and white flies.
10. Stages 7-9 are repeated until the crop is harvested or abandoned.
A parallel situation has been reported from Vietnam (ICRISAT, 1991a)
where farmers have been known to apply insecticide to groundnut every
day and still have crops that are completely defoliated by S. lilura.
We believe that this treadmill effect can be avoided at the very begin-
ning. Farmers apply insecticides because they want to protect their large
financial investment in groundnut seed from (cosmetic) jassid and thrip
damage. There is available a variety (ICGV 86031) that has high yield
potential along with resistance to jassids and thrips (and, therefore, bud
necrosis virus) and defoliators, as well as foliar disease. We are testing this
variety in key areas on the farms of lead farmers. Although our experience
Integrated control of ground nut pests 465
at ICRISAT indicates that pest outbreaks can be reduced by holding back
on insecticide application (Table 11.21), we see the need to give farmers
the confidence to get off the treadmill. We hope that the provision of a
new, suitably adapted variety will be accepted as an alternative to a heavy
insecticide application regime.
We are not as confident of being able to subdue the groundnut leaf miner
with host plant resistance at the same time as the other potential pests in
this environment. However, we have evidence that our action thresholds
for insecticide application for leaf miner control are realistic and can be
handled by farmers, and that the parasite cadre is sufficiently robust to
survive limited and selective insecticide application (Table 11.5).
Continuing research will cover the role of pheromone trapping in dealing
with this problem and the possibility of exploiting wild species genes.
This reflects the importance of developing varieties with multiple pest
constraint resistance. Even though there may be varieties with higher yield
potential, they may never achieve it on farm because pests and the farmers'
reaction to them would not allow it. Similarly, a single constraint approach
to pest management, followed to the exclusion of consideration of the
needs of the system as a whole, can also lead to sub-optimal solutions. For
instance, there is little logic or acknowledgement of the principles of IPM
to be found in developing a variety that has resistance to a single con-
straint, such as a virus disease, that hits seriously in a particular zone in
only one season in 10 or 20. It is more sensible to concentrate on multiple
insect resistant lines that reduce the risk of one-in-two season insect
outbreaks and give perhaps 70% virus control via vector resistance. We
have also indicated that a general need for groundnut is foliar disease
management, with insect resistance being required in specific agroecologi-
cal zones.
We agree with Lynch and Douce (1992) that the future of IPM for
groundnut crops will involve computer modelling. This will emphasize,
even more, the need for IPM researchers and practitioners to adopt
multidisciplinary approaches. For instance, those interested in pest
damage need to be able to discuss leaf eaters with physiologists in terms of
depleted leaf area indices and disturbed source-sink relationships in the
case of root feeders.
It will be noted that there is in this chapter data that form the basis of a
forecasting system for a defoliator (in this case S. litura) management.
Here are its components, with cross-references to appropriate sections.
1. Flight intensity of migrant moths is monitored by means of a district-
wide network of pheromone traps (section 1l.3.1(a)). Trap data are
reported daily to a central facility.
(Alternatively, scouts could visit fields to count eggs or egg masses.
This is not a preferred procedure because it gives farmers less lead time
to prepare to treat their fields during the first ins tar .)
466 Groundnut pests
2. Data for each trap or group of traps are entered into a software package
that, on the basis of those data and on a given day for a daily cohort, will
estimate:
(a) the density and date of first day fourth instar larvae and of the
moult from third instar to fourth instar based on current prediction
models and daily mean air temperatures for the district (sections
11.3.1(b) and 11.3.2(a));
(b) the total leaf area removed per day of that cohort during instars
IV-VI (from the energetics model in section 11.3.3(a));
(c) the effect of the insect-induced reduction in leaf area on pod
production (via a crop model running off the same meteorological
data as the insect popUlation density model) (section 11.3.3(b)).
(d) whether the data for the cohort of that day, combined with the
information from previous cohorts, will result in a yield reduction
caused by the cumulative activity of the IV-VI larvae (note that the
level of resistance to the defoliator in the various genotypes grow-
ing in a particular district would have to be taken i.nto account in
this stage and if antibiosis retards larval development in 2(a);
3. Output would be in the form of advice to farmers in the village or
watershed to apply insecticide, covering:
(a) the selection of materials to be applied;
(b) the preferred date;
(c) the rate, as well as linked advice about harvest dates, etc.
Certain strategies should be built into this third stage. The first is the
application of low rates of insecticides to conserve natural enemies, early in
the season. Ideally, farmers should apply insecticide just as the eggs hatch.
This is when the larvae are at their most susceptible to insecticides, so that
much lower rates than would be used against larger larvae can be applied.
This strategy is fundamental to the conservation of natural enemies.
However, it is not easy to achieve, because sharp eyes and constant
surveillance are required. Farmer participation would be needed to build
confidence in the timing of this procedure,which would be based on a
physiological constant.
We are aware that populations can explode later in the season if the
correct action is not taken early enough. This means that fall-back strate-
gies are required. We look to the exploitation of insect pathogens and
perhaps selected natural insecticides to supplement them.
We are also aware that the misuse of insecticides has resulted in the
development of chronic levels of insecticide resistance in developing
countries. This can be offset, with the co-operation of the agrochemical
industry, by a planned approach to the phasing of given insecticides and
the alternation or rotation of active ingredients to avoid or manage insecti-
cide resistance.
There is a final component in this management system:
Conclusions 467
4. The information should be returned to the farmers within hours by
whatever means are available: radio, television, telephone to village
leaders of 'pest management clubs' or, simply, a person on a bicycle.
This covers an approach to managing S. litura that could drastically reduce
the amount of insecticide applied in several parts of Asia, without endan-
gering the productivity of the farming systems concerned. The basic data is
available but needs integrating and verifying in farmers' fields. Others
crops that are susceptible to S. lilura, such as tobacco and vegetables, could
also be integrated. Virtually all of the relevant information is available for
the groundnut leaf miner which could be 'added on', perhaps even includ-
ing a function relating pheromone-trap catch data to larval densities once it
has been established. Soil insect management requires a different approach
involving soil sampling and will not be discussed here.
11.10 CONCLUSIONS
Throughout this chapter we have indicated where there is scope for further
research activity and where significant progress has been made. We have
distinguished the need to set two kinds of priority - research and economic
- on the grounds that specific insects can remain of great (potential)
economic importance, even though researchers have provided all the
information (within the bounds of the current technology) needed to
manage them. Clearly, if and as the fruits of research become accepted as
on-farm practice, the economic status of an insect should diminish: vide
Aphis craccivora.
This is why we have divided the priority ranking of pests in Table 11.24
between economic and research. This table, which is presented as a basis
for thought stimulation, challenge and discussion, as much as for infor-
mation, indicates the need to pay considerably more attention to the soil
zone than it has been given in the past. A comparison of Table 11.24 (here)
with Table 6 in Smith and Barfield (1982) indicates where progress has
been made - for instance, in our understanding of thrips and their relation-
ship with groundnut management - and where recent research has unco-
vered the need to put emphasis in new directions, for instance white grubs.
Since Feakin completed the third edition of her comprehensive hand-
book in 1973, there has been a considerable consolidation in our knowl-
edge of groundnut insects. It is clear that, in the USA, groundnut IPM has
passed from the research to the implementation and extension phase.
Looking at the American scene from a long distance, there appears to be a
need for integration across political boundaries and deciding where com-
mon goals have been achieved, where voids in knowledge appear and who
can best fill them.
In less developed countries, it is possible to detect all levels of progress.
TABLE 11.24 Major groundnut pests of the world with an indication of their ranking in terms of the need to carry out research on them and
their economic importance in the relevant agroeconomic zones together with an indication of current and potential management solutions
Importance Current solution Potential solution
Research Economic
Millipedes None ?
White grubs Insecticides (India) Resistant genes in Arachis spp.
None in Africa Slow release insecticide formulations
Metarrhyzium
Cultural control
Termites None in Africa Resistance to scarification
Insecticides in Asia
Hilda None ?
Pod borers (wireworms, false
wireworms, doryline ants, earwigs) 2 None ?
Elasmopalpus lignosellus 2 Granular insecticides
Irrigation
Diabrotica 2 2 Insecticides Host plant resistance
White fringed weevil 2 2 ? ?
Miridae 1 2 None ?
Thrips:
Brazil 2 1 Insecticides Host plant resistance
as TSWV/BNV vectors 3 2 Host plant resistance Host plant resistance
Spodoptera litura 1 2 Insecticides (excess) Host-plant resistance, especially via wild spp. genes
Natural and cultural control via IPM
S. frugiperda and S. exigua 2 2 IPM via pheromone traps
monitoring (USA)
Heliothis zea 2 2 Potential for natural control plus cultural control to
obviate need for insecticide
Groundnut leaf miner Insecticides Host plant resistance, especially via wild species genes
IPM, including pheromone trap monitoring
Hairy caterpillars 2 2 Insecticides and cultural practices ?
Aphis craccivora 3 3 Cultural Host plant resistance
Jassids 3 3 Insecticides (if anything) Host plant resistance
Ranking (scale of 1-5): 1= of greatest importance

References 469
Unfortunately, the pest spectrum and the farming systems (and their
economics) in the USA (and Latin America) are so different to those in
Africa and Asia that it is not easy to see direct ways of transferring suitably
modified technology. On the other hand, recognition that the high levels of
insect resistance in NC Ac 343, 2240, 2214, etc. (which were developed at
North Carolina State University from irradiated material) are effective
outside the USA and are heritable has given entomologists and breeders a
'flying start'. These and other more recently identified sources of host plant
resistance, and hopefully genes from the wild species, will surely form the
basis of the successful management of groundnut pests in the future.
ACKNOWLEDGEMENTS
Most of this chapter was compiled while J.A. Wightman was on sabbatical
leave and we thank Dr L.D. Swindale (Director General Emeritus) and Dr
J.G. Ryan (Director General) of ICRISAT for permitting the sabbatical
and also Dr J.G. Miller (Director General of the Queensland Department
of Primary Industries) and Professor Marcos Kogan (Director of the
Integrated Plant Protection Center, Oregon State University) for their
hospitality. Professors M.E. Irwin and Gail Kampmeier (Illinois Natural
History Survey) provided invaluable information, just as it was needed.
Special thanks go to Dr Robert Lynch (USDA, Georgia) for drawing our
attention to relevant sections of the literature and for providing un-
published information. We also thank Dr Ali Niazee (Oregon State
University) and Dr Lynch for their detailed comments on the manuscript.
We are aware that we have absorbed the thoughts and experience of
colleagues in many countries and embodied them perhaps rather tersely in
this manuscript: their anonymous contribution is duly acknowledged.
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CHAPTER 12
Industrial utilization and processing

].].K.B. Asiedu
12.1 UTILIZATION
The groundnut or peanut (Arachis hypogaea L.) is grown for its kernels,
the oil and meal derived from them, and the vegetative residue (haulms).
The kernels may be eaten raw, roasted or boiled, sometimes salted or
made into a paste popularly known as peanut (or groundnut) butter. The
tender leaves of the plant are used in certain parts of West Africa as a
vegetable in soups. In Senegal, one of the leading countries in the pro-
duction of groundnuts, roasting and selling of groundnuts are done by
women (Figure 12.1).
Groundnut oil is perhaps the most important product of the crop. At
present about 40% of the world crop is processed into oil, which has a
multitude of domestic and industrial applications. It may be used for
cooking, for margarines and vegetable ghee, for shortening in pastries and
bread, for pharmaceutical and cosmetic products, as a lubricant and emul-
sion for insecticides, and as a fuel for diesel engines (Duke, 1981). The
press cake containing 40-50% protein is used mainly as a poultry feed.
Groundnut flour, produced from the cake, can be used for enhancing or
enriching the nutritive value of tuber flours that are low in protein, such as
cassava flour. The so-called 'Mysore Flour', which is a combination of 25
parts of groundnut flour and 75 parts of cassava flour and contains a protein
content of 12% , is utilized as a partial substitute for cereals in India. Also
in India, 'Paushtik atta', a blend of wheat flour, groundnut flour and
cassava flour, is used for the preparation of chapattis - an unleavened
bread.
The dry pericarp of the mature pods (known variously as shells or husks)
may be used as fuel, as a soil conditioner, as a filler in feeds and as a source
of furfural, or processed as a substitute for cork or hardboard, or com-
posted with the aid of lignin-decomposing bacteria (Adams and Hartzog,
1980). The haulms of the groundnut crop are reported to be nutritionally
Utilization 481
Figure 12.1 Roasting and selling groundnuts in Dakar, Senega\.
comparable with grass hay and are used in many countries (Gibbons,
1980).
Groundnut or peanut butter is a comminuted food product prepared
from dry-roasted, clean, sound, mature groundnuts from which the seed
coat and germs (hearts) are removed, and to which salt, hydrogenated fat
and sometimes sugars, antioxidants and flavours are added. Peanut butter
is by far the most important product made from ground nuts in the United
States of America, consumption level reaching about 1.4 kg per person
(Zamula, 1985). While more than 90% of the groundnut butter is stabilized
with hydrogenated oil and antioxidants and consumed in homes or school
lunch rooms, the remainder is un stabilized and is used commercially in
confections, bakery goods, ice cream, breakfast cereals and in other minor
ways. Groundnut butter also enjoys considerable popularity outside the
United States. The countries in this respect include Canada, where con-
sumption level is 0.2 kg per person, The Netherlands, with a consumption
level of 0.16 kg per person and, to a lesser extent, the UK, France and
Germany (Wilson, 1975).
A substantial amount of groundnuts finds its way into candies and
confections. The characteristic flavour, crunchy texture and high protein
content render roasted groundnuts especially suitable for use in candies
and confectionery. Peanut candies and confections are considered to be the
482 Industrial utilization and processing
major growth area in groundnut consumption: 44.5% of chocolate bars and
3.7% of nut bars contain groundnuts (DEBS, 1984). Peanut candies and
confections encompass a wide variety of product types which include (but
are not limited to) nut roll candies, dragees, peanut brittles, peanut roll
bars and spun candy bars.
Groundnuts are consumed in large quantities in the salted form. Salted
groundnuts are shelled, roasted and salted to taste; they mayor may not be
blanched.
Among the products that have not yet obtained wide popularity are
groundnut protein, groundnut milk, groundnut cheese, frozen boiled
groundnuts, groundnut bread and partially defatted groundnuts.
12.2 PROCESSING
Processing is a unit operation or series of unit operations performed on a

raw material to change its form or composition. Processing can be as
simple as cleaning, grading and packing fresh produce or as sophisticated
as using, for example, pure strains of enzymes to convert maize starch to
high-fructose sweeteners. Traditional processing evolved from society's
need to enhance the usefulness of agricultural products. The basic methods
employed for this purpose are preservation and separation. Raw materials
can be converted to a form less susceptible to deterioration, usually
through drying, curing, fermentation or refrigeration. Some forms of
preservation, such as fermentation and cooking, impart other desirable
characteristics to the finished product that may become ends in themselves,
but they remain essentially preservation processes. Irradiation and freeze-
drying are the principal innovations in preservation during the past
century.
Desirable components such as seed, oil or fibre can be isolated from the
parent material occurring in nature. After the natural material has been
disaggregated, usually through the application of mechanical force, separ-
ation is achieved by exploiting differences in the weight, specific gravity,
size, shape, air resistance or solubility of the components. The use of
catalysts and solvents has been the principal development in this type of
processing in the past century.
Depending upon the desired product, groundnuts may be subjected to a
number of unit operations which may include picking or stripping, clean-
ing, storing, shelling, blanching, roasting, salting, grinding, extraction or
expression.
Harvesting, stripping and drying 483
12.3 HARVESTING, STRIPPING AND DRYING
12.3.1 Harvesting
The timing of the groundnut picking or harvest is critical since it can
significantly affect the yields and nut quality. Groundnuts are best har-
vested or picked when the soil is dry enough to fall freely from the stems
and pods. However, if the soil is very dry and crusted many of the pods
may break off and remain in the soil. In ~ubsistence agriculture, harvesting
consists of either digging or pulling up the plant manually from the soil,
allowing the plant to wilt and then picking the pods from thc vines. On
large-scale farms, groundnuts are harvested by running a special groundnut
digger or wing-type plough below the surface and under the plants.
Commercial peanut combines are also available. These consist essen-
tially of a thresher, a pick-up feeder chute and an auxiliary engine. The
pick-up feeder chute is attached to the front, while a long bagging elevator
and a high bagging platform are fixed at the rear. The machine is powered
by an auxiliary engine. The speed of the pick-up cylinder in such combines
approximately equals the forward travel of the combine. The following
types of combine harvesters are generally in use on commercial farms.
- Self propelled
- Tractor-drawn, p.t.o. driven
- Tractor-drawn, auxiliary engine driven
- Mounted, auxiliary engine driven.
12.3.2 Stripping
The process of separating groundnut pods from the plant or haulm is
known as stripping, which may be accomplished manually or using mech-
anized means. Manual stripping, that is picking pod by pod, is time and
labour consuming although this operation results in good physical appear-
ance of the groundnuts (fewer pods with vine attached, and clean). The
stripping capacity of such a practice varies from 2 to 6 kg/man-hour. In
some areas farmers strip groundnuts by hitting or beating a handful of the
harvested plants against a stick placed on top of a container. This method
results in much higher stripping capacities - from S to about IS/kg
man-hour, depending upon the moisture content of the plant. The major
drawbacks associated with this method are high scattering loss and a
substantial amount of impurities.
Groundnuts may be stripped in a Drum Stripper, in which a handful of
groundnut plants are beaten over the rubber-covered rods of a revolving
drum. The output of this stripper is about 16 kg/man-hour.
Over the last decade a number of other designs of stripper have been
developed and tested and found appropriate for small-scale and medium-
sized farms. These include:
- Engine-operated spike-tooth stripper consisting mainly of a rotating
cylinder provided with appropriately spaced spike teeth and a concave.
- Hold-on mechanical stripper in which groundnut plants are held against
stripping bars of a rotating cylinder until pods are detached.
- Manually operated paddle stripper/winnower. This unit consists of a
stripping cylinder and a drive mechanism (Figure 12.2). Stripping is done
by operating a foot on the paddle so that the stripping cylinder, which
has eight steel rods as stripping bars, rotates outwards from the
operator.
- Modified axial flow rice thresher. This consists of an upper concave with
fins, and a rotating drum. Material fed in at one end of the drum passes
along the drum as it is rotated, and is ejected at the other end. Stripping
occurs as the groundnut plant passes between a revolving cylinder and a
metal grate (concave), which covers part of the circumference of the
drum.
12.3.3 Drying
At the time of digging, groundnuts generally contain 35-60% moisture.
Until the moisture is reduced to below 10%, the nuts are prone to mould
attack, especially at warm temperatures and high humidity. Groundnuts
can be dried to about 8% in field stacks in 4-8 weeks and to 9% in
windrows in 1-2 weeks. Mechanical driers using heated air may be used.
Lid
Figure 12.2 Paddle stripper/winnower (Khon Kaen University/IDRC, 1987).

Storage 485
The air should not be heated above 44 DC for groundnuts which are meant
for table stock; temperatures up to about 52 DC can be used if the nuts are
to be used for oil. On a continuous drier, with the groundnuts on the vines
placed 0.6 m deep, the best drying and quality are obtained by using an air
velocity of 0.30 mls at 30-32 DC. The major problem in drying groundnuts is
to prevent the splitting of the skin caused by rapid drying.
12.4 STORAGE
Groundnuts are known to be semi-perishabl~, which means that, on the

one hand, they may be held for a number of years under optimum
conditions. On the other hand, under unfavourable conditions they be-
come inedible within a month due to mould, insects, absorption of foreign
flavours or rancidity. For successful storage, the following requirements or
conditions should be met as far as possible.
The storage temperature should be low. In general, the lower the tem-
perature, the longer the expected storage life of groundnuts, although no
critical temperature has been established at which groundnuts must be
held. Table 12.1 shows the relationship between temperature and the
length of time the nuts retain their edible quality. At about 21 DC shelled
groundnuts are susceptible to insect infestation, development of an 'amber'
colour, staling and rancidity. At about 8 DC insect development is arrested
and shelled nuts may be held for 6 months.
TABLE 12.1 Relationship between temperature and

storage time of groundnuts
Time of retaining edible quality
Temperature (0C) In-shell Shelled

groundnut groundnuts
21 6 months 4 months
8.3 9 months 6 months
0 3 years 2 years
-4 7-8 years 5 years
-12 15 years 10 years
The relative humidity should be between 65% and 70% . High moisture in
groundnuts is reported as being possibly the cause of more deterioration
than any other single factor. Above 70% humidity, groundnuts are likely
to develop mould. Below 65%, the nuts lose weight, become brittle and
may split during handling.
The atmosphere in the store should be free of odours and flavours and
well aerated. Loss of natural flavour of groundnuts increases with temp-
erature and, likewise, absorption of foreign flavour increases with
temperature.
12.5 SHELLING OR DECORTICATING
The terms 'shelling' or 'decorticating' are used in groundnut processing to

describe the separation of the kernels or seeds from their shells or husks
through the application of mechanical forces. In most cases compressive
forces (sometimes, however, in combination with shear forces) are
required to crush open the pod so that the kernels can be freed from the
shells.
Shelling is usually carried out for two reasons:
1. Groundnuts in the shell are about 50% heavier than kernels alone and
are therefore costlier to transport.
2. Groundnuts have to be shelled to facilitate further processing.
Shelling has the disadvantage that kernels removed from pods are more
susceptible to attack by insects and moulds than in-shell kernels.
Smallholder farmers shell their groundnuts manually, using finger press-
ure. This is an arduous, boring, labour-intensive and low output operation
which, in several cases, leads to the so-called 'thumb-sore syndrome'. The
unique advantage of this method is its excellent outcome as regards
cleanliness and minimum breakage and wastage of the shelled kernels.
Waste is almost eliminated, and intactness and cleanliness of the kernels
have great effect on the choice of groundnuts for seeds.
A number of groundnut shellers are available for various levels of
production and powered by various means (manual, diesel engines, electric
motors). Splitting or breakage of kernels can be as high as 22% during the
operation of the more efficient shellers or decorticators. To reduce splitting
and breakage to a minimum, groundnuts are shelled at 8% moisture (wet
basis) or higher.
12.5.1 Hand-operated shellers

The basic design for hand-operated equipment is relatively simple and
common to all makes. Essentially, the devices consist of a semi-cylindrical
screen, a handle, a shelling grid which forms part of the outer casing, and
spiked cast-iron rustlers attached to the handle. The action of the rustlers is
effected by moving the handle back and forth. This causes the nuts to be
The processing of groundnuts into peanut butter 487
rubbed between the rustlers and the shelling grid. The shelled kernels and
crushed shells fall through the perforations of the shelling grid into a
receptacle placed below it. This mixture of kernels and shells may be
separated using air suction or a winnower.
The 'sliding' groundnut sheller is another hand-operated device consist-
ing of a base, hopper box, adjustable throat slide, removable grid and
decorticating bars. The box forming the hopper and accommodating the
decorticating bars is pushed back and forth, using handles at both sides,
above a removable grid. Shelling takes place in the clearance between the
decorticating bars and the grid. The brok'en shells and the freed kernels fall
through the gaps of the grid into a container placed below (Asiedu, 1987).
One of the most recently developed manually operated machines is the
rubber tyre groundnut sheller (Khon Kaen University Report, 1987). This
consists of a rubber tyre assembly, a concave, a hopper and a trough, with
a crank or handle for the rotation of the tyre. The shelling takes place in
the space between the tyre and the concave. Shelling capacity is reported
to be 60-80 kg (pod) per hour, with a shelling efficiency of 95%.
12.5.2 Power decorticators/shellers

The basic elements of most of the power groundnut shellers are frame-
work, drive shafts and pulley, hopper, electric motor or diesel engine and a
shelling chamber, where the groundnuts are shelled by rotating flexible
beaters. A fan, usually fixed below the wire-mesh chute, blows the shells
upwards, and the kernels are recovered tbrough a spout at the bottom of
the machine. Depending upon the model, the hourly output of such
decorticators varies from 150 kg to 400 kg.
A motorized version of the manual rubber tyre sheller enjoys consider-
able popularity among large-scale farmers in Thailand (Figure 12.3). The
sheller is powered by a 1.49 kW electric motor and fitted with a blower and
a three-layer sieve. It has a shelling capacity of 300 kg (pod) per hour and a
shelling efficiency of 95%. However, its use is not recommended in shelling
groundnuts for seed (Khon Kaen University Report, 1987).
12.6 THE PROCESSING OF GROUNDNUTS INTO PEANUT

BUTTER
Peanut butter is a cohesive, comminuted and palatable food product

prepared from dry-roasted, clean, sound (in terms of maturity and attack
from insects and moulds) groundnuts from which the seed coats or skins
and germs are removed and to which salt, hydrogenated fat and sugars,
antioxidants and flavours are added. Although the manufacture of peanut
butter appears to be relatively simple, a complex interaction occurs be-
tween the components. Essentially, the manufacture of peanut butter
Figure 12.3 Motorized rubber tyre sheller (Khon Kaen University/IDRC, 1987).
consists of shelling, sorting, dry-roasting and blanching the nuts, followed

by two stages of fine grinding and packing in hermetically sealed glass or tin
containers. Ingredients such as salt, hydrogenated fat and dextrose, gly-
cerin and lecithin or antioxidunts are usually incorporated to improve
flavour, prevent oil separation and control rancidity. A simplified flow
chart for the processing of peanut butter is shown in Figure 12.4.
12.6.1 Processing and equipment

Shelled groundnuts are sorted to remove discoloured kernels and other
undesirable contaminants before they are roasted. Roasting is done either
on a conveyor belt passing through heated air, or in a rotary roaster.
During roasting the kernels are heated to about 160-170 °C and are held at
this temperature for 40-60 minutes to achieve the desired level of roast.
After roasting the kernels are transferred to a perforated cylinder or cooler
vat, where air is blown through the mass by fans.
Cooling is followed by dry blanching in a rotary cylinder or blancher.
Blanching consists of removing the skins (seed coats) and the hearts of the
kernels, which would otherwise give a bitter and rancid-like butter, by
exposure to 126-145°C heat for 5-20 minutes, followed by rubbing the
kernels between soft surfaces and removing the skins by blowers and
the hearts by screens. The blanched kernels are inspected and screened,
and scorched or rotten ones are discarded.
The nuts are then ground, in one or two stages. In the first, they are
Defects Hearts (germs)
Discoloured kernels Skins (seed coats)
Blanching
I - - -...
·~I Sorting I·[ Dry-ro:ng-- • l...-.-------r------'
Mixing, degassing Second stage Addition First stage

and cooling grinding of ingredients grinding
Packaging
Figure 12.4 Manufacture of peanut butter.

ground into butter by attrItion mills, homogenizers, hammer mills or
colloidal mills. Ingredients like salt, hydrogenated oil, dextrose, corn syrup
or honey and stabilizers are then added to improve smoothness, spreada-
bility and flavour. A second, fine-grinding stage may follow and this fine-
ground butter is mixed, degassed, and cooled in a rotating refrigerated
cylinder before being packed, labelled and stored. Peanut butter contains
about 1-2% moisture, 49-52% fat, 27-29% protein, 5-17% carbo-
hydrates,4% phosphorus and 2% fibre.
12.6.2 The role of ingredients and additives

A food additive is defined as:
a substance or mixture of substances, other than a basic foodstuff,
which is present in food as a result of any aspect of production,
processing, storage and packaging. The term does not include chance
contaminants. An additive may be either nutritive or non-nutritive; it
may be physiologically active or inert; it may be present intentionally,
to achieve some modification in the foods, or incidentally and serving
no useful purpose in the final product
(National Research Council of the Food Protection
Committee of the National Academy of Sciences, 1959)
In general, there are seven major reasons for the use of additives: taste,
appearance, texture and eating quality, processing, stability or preser-
vation, nutritional benefit, and for special and dietary foods.
In particular, processing additives make possible the use of many in-
gredients by the food processor and the consumer not otherwise available
at the desired quality level. For example, without the use of process
additives such as monoglycerides, instant mashed potatoes would have the
qualities of a sticky paste.
Before World War II, peanut butter is reported to have been prepared
from groundnuts, salt and sugar 'only'. This resulted in oil separation,
remixing difficulties, stickiness, graininess, poor spreadability, rancidity
and poor shelf life. The addition of stabilizers prevented these defects and
provided transportation and storage stabilities, and manufacturers have
continued to rely on various stabilizers since 1945.
Glycerin and monoglycerides prepared from vegetable and groundnut
oil are among stabilizers which prevent oil separation. Stabilizer selection
generally depends upon the process and desired flavour release, grind,
packing season, and fill and storage temperatures. Fine grinds produce
more 'free' oil and require more stabilizer (0.1-0.15% or more). Single-
stage grinds require a different stabilizer than two-stage grinds since the
product leaves the mill at a higher temperature. Hard butters, which set
rapidly after filling and cooling, can result when excessive stabilizer is used.
At low fill temperatures of 35-44°C, the stabilizer rapeseed oil hardfat
The processing of roasted and salted groundnuts 491
(1.6-1.8%) is preferred for summer packs and results in less 'pull away'.
For higher fill temperatures (49-54 0c), followed by a chill tunnel, cotton-
seed hardfats (1.8-2.0%) are used. At even higher fill temperatures,
monoglyceride stabilizers can be used along with glycerol monostearate
(Weiss, 1983). Use of stabilizers can be avoided by refrigerator cooling (at
less than 10 0c) or by adding absorbants like ground groundnuts, powdered
sucrose, dried milk or partially defatted wheat germ (Woodroof et ai.,
1945). Cooling and storing peanut butter at less than 10 °C, however, lead
to increased storage and energy costs. ,
The choice of a sweetener depends upon the process. Maillard reaction
between amino acids and reducing sugars can be accelerated by high
processing temperatures to give dark peanut butters with altered flavour
and texture. Sweetness and a smooth, non-sticky mouthfeel can be ob-
tained with corn syrup solids (28-48% reducing sugars), while molasses
imparts a unique flavour and consistency, although the high moisture
content of honey (21 %) favours lipid oxidation and rancidity. Also, honey
or corn syrup, due to their high moisture, can lead to butters with high
viscosity or pastiness (Weiss, 1983). The problems of lipid oxidation and
the development of rancidity can be avoided using lecithin fatty acids or
fatty acid polyglycerol esters at 0.15-0.85% of the weight of the honey.
Lipid oxidation and rancidity can also be delayed by storing peanut butter
in nitrogen atmospheres. Stickiness in peanut butter is reduced by adding
partially hydrogenated vegetable oils.
12.7 THE PROCESSING OF ROASTED AND SALTED

GROUNDNUTS
12.7.1 Roasted groundnuts

The unit operations involved in the making of roasted groundnuts (Figure
12.5) include roasting by heated air or oil cooking, blanching, sorting,
salting/glazing and packaging. Shelled groundnuts are passed through
cleaning screens to remove contaminants and other undesirables, separ-
ated by gravity tables or cyclones. After they have been white roasted (see
below), the nuts are put into colour sorters or 'electric eyes' where dis-
coloured and damaged nuts are rejected. The acceptable nuts are fed into
blanchers, where the red skins are removed but (usually) not the germs.
Dust, mould and other foreign material are also removed during blanch-
ing. The blanched nuts are again colour sorted, to discard unblanched and
damaged ones, and then roasted.
The first effect of roasting by heated air is rapid drying during which the
moisture content is reduced to about 0.5%. This is followed by the
development of oily translucent spots on the surface of the cotyledons.
Packaging Salting Cooking Frying
Cleaning Separating by
White Colour Colour
gravity table Blanching
by screens roasting sorting sorting
or cyclone
'Undesirables' 'Undesirables' Discoloured and Skins Unblanched

damaged and damaged
groundnuts nuts
paCkagingr-1 . ·~--r-I Coating r-I Cooling _---l

r!,----! Roasting
Figure 12.5 Processing of salted groundnuts and dry roasted groundnuts.

The processing of roasted and salted groundnuts 493
These 'steam blisters' are caused by the oozing of oil from the cytoplasm as
free oil. Changes in nut colour, due to cell walls becoming wet with oil, are
observed. This stage is usually referred to as white roast. The final stage of
roasting is brown roast: the nuts develop a brown colour and roasting is
completed.
Roasting is followed by cooling. In many industrial plants today the hot
nuts emerging from the roasters are conveyed directly to a perforated
cylinder or cooler box where fans blow air through the mass of the hot
nuts.
After cooling, the nuts are coated. Coatings used include heated
sorbitol-mannitol (7:3) blends or 5-6% acetic acid activated wheat glutins
(Gutcho, 1973).
Coating is followed by salting. Electrostatically deposited flour salt of
70-200 mesh is reported to be preferable for air-roasted nuts (Matz, 1976).
The type of salt applied in the processing of both roasted groundnuts and
salted groundnuts is of considerable importance as regards rancidity and
flavour. Groundnuts are rich in oils and since copper and iron (which are
important impurities of salt) favour and accelerate rancidity of oils and
fats, it is essential that a salt with minimal copper and iron impurities is
used. Non-refined sea salt high in CaCh is also reported to contribute to a
harsh bitter flavour (Woodroof, 1983).
Finally, the roasted groundnuts are packed in airtight containers to
retain 'freshness'. Methods include vacuum packing in cans and glass jars
and the use of nitrogen-flushed laminated foil packs.
12.7.2 Salted groundnuts

Operations required for the processing of salted groundnuts are essentially
the same as those for roasted groundnuts (Figure 12.5). The most con-
spicuous difference between the two products is the type of roasting they
undergo. Salted groundnuts usually undergo oil-roasting or frying. This
involves immersing the blanched and undamaged nuts for a period of 3-5
minutes in heated oil (coconut, groundnut or cottonseed oil) using a basket
or submerged belt, or frying in solid fats (for instance, coconut oil). When
nuts are fried in solid shortening, salt is applied before the nuts have cooled
to room temperature (24-25 Qe). Weiss (1933) reports that silicones
(0.5-3 ppm) are often added to the heat transfer medium (coconut,
groundnut or cottonseed oil) to raise the smoke point and reduce foaming.
Cooking oil should be changed frequently to prevent rancidity. Rancidity
can be detected within 30 days at 50% relative humidity (RH) at 24-28 QC
for groundnuts cooked in used oil, compared with 60 days for those heated
in fresh oil.
Salted groundnuts must be kept at less than 4% moisture, with 2-2.5%
moisture giving the best texture, flavour and appearance retention. They
will remain fresh for up to 35 days at 50% RH but storage at 80% RH leads
to a loss in freshness within 15 days (Woodroof et ai., 1945).
12.8 THE PROCESSING OF PEANUT CANDIES AND

CONFECTIONS
The use of roasted groundnuts in making different types of candies and

confections is due largely to their pronounced flavour, crunchy texture and
high protein content.
Unlike groundnut butter and roasted groundnuts, where the product has
a more or less uniform formula and processing technique, peanut confec-
tions encompass a wide spectrum of product types and processing tech-
niques. However, the basic unit operations used in making peanut butter
and roasted peanuts still find their application in the processing of peanut
candies and confection. Figures 12.6 and 12.7 demonstrate flow charts for
making the different types commonly found on the market.
More than 50 types of candy are made from groundnuts - possibly the
most popular ingredient in American candies. For the preparation of
peanut brittles, a mixture of sugar, corn syrup and water (30:20:7) is boiled
and groundnuts are added until the mixture reaches 154-157°C. Sodium
bicarbonate is added to aerate the brittle. The resulting batch is cooled to
about 3°C prior to packaging. It is reported that the cooking of the nuts
within the batch gives improved flavour, through retention of volatile
flavours, but the high moisture content of the sugar solutions encourages
browning, development ofrancidity and reduced shelflife (Janssen, 1978).
In dragees and hard-coated peanuts. the nut or a suitable peanut butter
formulation is coated with alternating supersaturated sugar/gum arabic
solutions (67%) and powdered granulated sugar. The final coating uses a
51 % sucrose syrup, which is dried at a temperature of 37-46°C before
packaging (Minifie, 1964). For the preparation of nut roll candies, ground-
nuts as the 'raw material' form a part of the nougat formula or are part of
the topping. Nougats are prepared by mixing sugar-corn syrup, egg white
and fat. This mixture is then whipped, extruded and finally coated with
chocolate.
Peanut candies and confections are relatively high in fats which tend to
shorten the shelf life. Stabilities differ, depending upon formula.
According to Labuza (1982), a typical shelf life is one year with distribution
times of 20-60 days. Uncoated peanuts - peanut brittles and peanut roll
bars - are more prone to rancidity. Brittles with 1.6% moisture were
reported to be stable for 1-4 weeks at 40-45% RH, while peanut rolls
having 5.89% moisture retained their original characteristics at 50-
55% RH. Stickiness is controlled by low humidity, while staleness is
checked by low temperature, i.e. below O°c. High temperatures (above
26°C) and humidities over 50% lead to sticky, dark and runny candies.
Addition of egg
white and fat
A) Enrobing
Whipping Extrusion with chocolate
Peanuts are used as

coating, or are incorporated
B)
Boiling EIrIttIe: Cooling
Addition of Addition of sodium

raw nuts bicarbonate
Packaging
Sugar/gum arabic
solutions
C)
Coating Drying Packaging
Figure 12.6 Making of different peanut candies and confections.

Addition of syrup, Nuts Sugar/gum arabic solutions
egg w\te and fat sy'v Nuts (initial coating)
Nougat ] I --...:B_ri_ttl_e_
/
Dragees
Groundl'll..lt$
Defects or
Separating by undesirables
gravity table or
cyclone
WhiteUr~sting~[ Coloursorting ~[ Bla~Ching~[ Colour sorting }------.-I brown roasting
I
Defects
I
Skins
I
Un blanched and
defect nuts
Chocolate
nut enrobing
Sucrose-syrup
'coated'
dragees
Figure 12.7 Making of candy nuts and dragees.

Oil expression 497
12.9 OIL EXPRESSION
Expression is the separation of liquids from solids by the application of

mainly compressive forces. Three methods of expressing the liquid from
the solid-liquid matrix are employed: hydraulic, roller pressing and screw
pressing. Hydraulic and screw presses are widely used in processing fruit
juice and oil seed; roller presses are universally used in expressing juices
from sugar cane.
The efficiency of an expression process depends on several factors,
including:
- the yield stress of the solid phase, i.e. its resistance to deformation;
- the porosity of the cake or mass formed during expression;
- the viscosity of the liquid expressed;
- the compressive force applied;
- the moisture content of the material to be expressed;
- material preparation or treatment prior to pressing.
In his study of factors affecting the yield of a number of vegetable oils, Koo
(1942) developed the following general equation:
W = C*WO*pI/2*t1l6*v-Z/2
where W is the oil yield (in weight %), C is a constant for the type of oil
seed (units consistent with unit analysis), Wo is the oil content of the seed
(in weight %), P is the pressure (in MPa), t is the pressing time (in hours),
v is the kinematic viscosity of the oil at press temperature (in m2/s) and Z is
the exponent of kinematic viscosity varying from % to liz. For groundnuts:
C, Wo and Z values are respectively 19.4*103 ; 51.9; 113. For anyone
oilseed there is an optimum range of moisture content for maximum oil
yield, from 5% to 13% (dry basis).
12.9.1 Processing groundnut for oil

The groundnut kernel is composed of approximately equal weights of fatty
and non-fatty constituents, the relative amount of each depending upon
variety and maturity. Most of the fatty constituents are found in the
cotyledons, which make up the bulk of the kernel or seed. All cells of the
groundnut embryo contain oil in an extremely fine emulsion. When kernels
are bruised or broken, a sufficient number of cells are injured and, as a
result, tiny drops of 'free' oil ooze out and collect on the surface of the
kernels.
Oil may be obtained from groundnut kernels through mechanical press-
ing (expression) or by the use of a solvent. Mechanical pressing makes use
of various types of equipment such as mortar and pestle, grinding stones,
hydraulic presses, expellers or screw presses, crushing rolls and rotary
ghanis. The rotary ghanis enjoy considerable popularity in the rural areas
of India.
Solvent extraction using hexane or cyclohexane is capable of recovering
nearly all the available oil from oilseed meal or flakes. In addition to the
high degree of oil recovery, the solvent method produces oil with better
qualities, and a higher protein meal. This method generally requires more
capital investment than a mechanical process. Mechanical pressing or
expression, though not as efficient as solvent extraction in terms of oil
recovery, has the advantage of producing end-products (oil and meal or
cake) free from dissolved chemicals and is a comparatively safer and less
costly process, demanding less skill.
Among the parameters which influence oil yield and oil quality are
pretreatment of groundnuts prior to pressing or extraction, operating
temperature, pressing time and moisture content during expression. Post-
extraction treatment of oil consists of refining and packaging.
(a) Hydraulic pressing

The use of hydraulic presses for oil extraction predominated until about
1945, since when the use of expellers has increased. For hydraulic pressing,
shelled and crushed groundnuts with an optimum moisture content of 6%
are heated by steam and then spread on press cloths, with the edges folded
to prevent running. The containers holding the heated material are
arranged one above the other (Figure 12.8). The racks are placed in tiers
about 1 m high. When pressure is applied, oil is released and is collected
beneath the tiers. As a press aid, a small amount of unshelled groundnut
meal is mixed with the crushed mass prior to pressing. The press cake
usually contains about 7% oil.
For expeller pressing, groundnuts are appropriately reduced in size and
fed into an expeller, which is basically a screw rotating within a cylinder.
The operation begins when the crushed ground nut mass is fed through a
hopper into the larger end of the expeller chamber and pressure is exerted
as the screw turns, forcing the mass towards the smaller or discharge end.
Friction and pressure cause the mass to heat, which facilitates oil extrac-
tion. The groundnut oil passes through the perforated screen walls and is
collected beneath the expeller chamber, while the press cake (which
normally contains 5% oil) is extruded from the discharge end or cake
outlet. The optimum moisture content of the groundnut mass for oil
recovery by expeller is about 6%. The main parts of an expeller are shown
in Figure 12.9. Expellers may be driven by an electric motor or via a pulley
and V-belt from a separate diesel engine.
The types in use are single, duo, or duplex expellers. A single expeller is
capable of pressing the assumed oil yield in a single pass. Such equipment
may have a capacity of 45-55 kg per hour, a power requirement of 3 hp and
Oil expression 499
Frame with press cylinder
Frame
Plunger
Press packet
Intermediate layers
Press floor
Figure 12.8 Multi-layer hydraulic press (Ulrich, 1967).
Hand-wheel
for adjusting choke
Ring
Drive shaft
Choke
Cake bars
Oil receiver
Figure 12.9 Oil expeller.

a rate of 300 revolutions per minute. In a duo expeller a 'first-pass' low
pressure and a 'second-pass' high pressure expeller are combined to
accomplish the extraction of the oil from the previously crushed ground-
nut. In a duo expeller the cake from the first pass is automatically conveyed
to the second. A duplex expeller has screws of varying pitch and differently
shaped chambers. This design enables pressure to be increased as the
material is moved through the chamber. Duplex expellers normally
achieve higher yields and faster throughput in a single operation.
In order to prevent the formation of groundnut 'butter' during extraction
of oil in small expellers, groundnut shells may be added to the groundnut
kernels.
(b) Solvent extraction

Oil extraction using expellers or hydraulic presses is always associated with
(or stigmatized because of) losses which can be as high as 10% or more; a
certain amount of oil always remains in the press cake, even with the most
efficient expeller. With solvent extraction it is possible to reduce the
residual oil in cake to less than 1%. The solvent extraction method for
groundnut is similar to that for soya bean. Generally, groundnuts for
extraction are shelled and winnowed to remove the fibre-rich shells, and
whitened by removing the tannin-containing red skins. The nuts are
cracked into pieces, conditioned to about 11 % moisture at a temperature
of 70°C and then rolled to form flakes, which may be cooked before they
are put into the extractor. Here the oil is extracted by means of a solvent.
The solvent-laden flakes are passed through a desolventizer to recover the
solvent. The extracted oil is clarified in a filter or frame press, after which it
may be dehydrated.
Solvent extraction plants may be divided into two main groups: batch
extraction plants and continuous extraction plants. The basic types of
equipment are open percolation tanks, extraction batteries and continuous
moving-bed extractors. Open mixing tanks, packed or plate columns and
centrifuge contactors such as Podbielniak machines are used for continu-
ous extraction. Percolation tanks are vessels with bottom outlets and
valves: the solvent percolates through the solids by force of gravity.
Percolation extraction batteries are tanks or vessels joined in series. They
have the same design features (bottom outlet, valve, etc.) as the percola-
tion tanks. The control valves are arranged so that a given vessel can be
separated from the others in order to be emptied or refilled. In all continu-
ous extractors the raw material moves through the vessels continuously
against the flow of the solvent. Continuous extractors vary in design but the
tower type is representative. This is equipped with travelling elements that
contain the solids, sprays to deliver the solvent to the solids, and a feeder
and discharger for the solids.
Oil expression 501
Degumming
'Crude Oil' Bleaching
Neutralization
L-_D_i_st,ill_at_io_n_----'~ Deodorization Filtration
Winterization
(for salad oil)
Hydrogenation
(for shortening and margarine)
Figure 12.10 Sequence of operations in the refining process of groundnut oil.
12.9.2 Post-extraction/pressing treatment

Most vegetable oils can be consumed in their unrefined state, after simple
mechanical pressing of the seed or fruit. But the colour and aroma of raw
oils mask other food ingredients, and more sophisticated markets demand
oils that do not dominate the sensory characteristics of food preparations.
Simple refining consisting of filtration and water removal gives a product
that has more acceptable cooking and shelf-life characteristics, but heat
must be applied at a fairly specific temperature and a filtration process
must be introduced. Today, however, most markets other than those in
isolated rural areas demand oil that has been fully refined, through bleach-
ing, deodorizing and the removal of free fatty acids to prolong shelf life.
These processes cannot be accomplished without precise control of the
temperature and material flow, and the use of caustic soda and bleaching
and deodorizing compounds that must be thoroughly separated from the
refined oil in additional stages of processing.
Groundnut oil obtained using a hydraulic press or expeller may be
refined for various reasons as elucidated above. Refining (Figure 12.10)
consists of alkali treatment, to neutralize the free fatty acids, using sodium
hydroxide; bleaching, to improve flavour stability, using natural bleaching
earth; and deodorizing, to remove odour, using steam distillation under
vacuum. If ground nut oil is to be used as a salad oil, it undergoes the
process of winterization prior to deodorization. Winterization is the re-
moval of fats that crystallize out at about O°c.
For margarine-base oils and shortening, groundnut oil is subjected to
hydrogenation, the process by which elemental hydrogen combines cataly-
tically with unsaturated organic compounds. It is conducted extensively in
the edible fats and oils industry to modify the characteristics of natural fats
and oils. Hydrogenation raises the melting point and increases the hard-
ness, so that it is useful in changing vegetable oils from liquids to solids
suitable for shortening and margarine.
Most hydrogenation is accomplished in batches, although equipment for
continuous operation is also available. The typical hydrogenator is a
carbon steel vertical pressure vessel. It is equipped with coils for heating
with steam and for cooling with water, turbine type agitators, stationary
baffles and a sparge pipe for admission of hydrogen. In operation, the
vessel is filled to its operating level with the oil to be processed. Usually the
feed has been refined and bleached and is free from water and dissolved
gases. The oil is first heated to approximately 93°C and a small amount
removed to a catalyst mix tank, where catalyst is added. The oil and
catalyst are then returned to the hydrogenator; where heating is continued
to 121-135 °C, at which point hydrogen is admitted through the sparge pipe
at a rate to maintain a predetermined pressure. The commonly used
catalyst is finely divided metallic nickel.
The degree of saturation of fats is determined by their iodine value. The
higher the iodine value, the greater the degree of unsaturation. The
common edible vegetable oils have iodine values ranging from 100 to 150.
Fats for shortening and margarine are hydrogenated to an iodine value of
70-90. These products are solid at ambient temperatures.
Groundnut oil is neutralized to dispose of free fatty acids which, when
oxidized, would cause rancidity. Neutralization involves combining the
free fatty acids with an alkali, usually caustic soda, to form a soap solution
which can be separated from the oil. After the soap has been separated,
the oil is washed and dried under vacuum. Then it is bleached with the aid
of a specially treated form of fuller's earth, which is capable of absorbing
colour pigments and residual oxidized material that would later affect the
stability of the oil. After bleaching, the oil is passed through a filter press
for removal of the fuller's earth and any other impurities, and is subjected
to deodorization, a process of steam distillation under vacuum. High-
pressure steam is blown through the oil so that the odoriferous materials,
which are steam volatile, concentrate in the distillate.
12.9.3 Filtration
Filtration is a unit operation designed to separate, by means of a porous
medium or screen, solids from liquids or liquids from liquids. Filtration is
involved in many branches of the food industry, such as the making of fruit
juices, vegetable oils, alcohol, sugar and beverages.
Filtration is accomplished by a filter medium through which a slurry is
Oil expression 503
separated into solids and liquid by means of gravity, pressure, or vacuum.
In the food industry the liquid is normally the valued or desired constituent
and the solids are the undesirables.
For the filtration of ground nut oil, a filter press is generally used. It is
made in two main forms: the plate and frame press, and the recessed plate
or chamber press (Figure 12.11).
The plate and frame press consists of plates and frames arranged alter-
nately and supported on a pair of rails. The plate has a ribbed surface and
the edge stands slightly higher. The hollow frame is separated from the
plate by the filter cloth, and the press is closed either by means of a hand
screw or hydraulically; minimum pressure should be used in order to
reduce wear on the cloths. A chamber is in this way formed between each
pair of successive plates, (Figure 12.12). The oil to be filtered or clarified is
introduced through a port in each frame and the filtrate passes through the
cloth on each side so that two cakes are formed simultaneously in each
chamber, and these join when the frame is full. The filtered oil drains into
passages at the base of the plate and from there it is collected. In many
filter presses, provision is made for steam heating so that the viscosity of
the filtrate is reduced and a higher rate of filtration obtained.
The chamber press is similar to the plate and frame type, except that the
use of frames is obviated by recessing the ribbed surface of the plates so
that the individual filter chambers are formed between successive plates. In
this type of press, therefore, the thickness of the cake cannot be varied.
The feed channel usually differs from that used on the plate and frame
press. All the chambers are connected by means of a hole in the centre of
each plate and the cloths are secured in position by means of screwed
unions. Slurries containing relatively large solid particles can readily be
handled in this type of press without fear of blocking the feed channels.
For filtering vegetable oils, the filter press has several advantages:
- Owing to its simplicity, it is versatile and may be used for a wide range of
materials under varying operating conditions of cake thickness and
pressure.
- Maintenance is low.
- It provides a large filtering area on a small floor space.
- Most joints are external and leakage is easily detected.
- High pressures are easily obtained.
- It is equally suitable whether the cake or the liquid is the main or the
desired product.
The disadvantages of the filter press include the following:
- It is intermittent in operation and continual dismantling is apt to cause

high wear on the filter cloths.
- Labour requirements are high and the throughput is moderate.
a1 d a b e c
a Filter elements e Press frame

a 1 Tail plate f Stand
a2 Head plate
@»~ ~ 9 Inlet
b Screw h Outlet
c Hand wheel Filter cloths
d Tension rod
Figure 12.11 Filter press (a, filter elements; ab tail plate; a2, head plate; b, screw; c, hand wheel; d, tension rod; e,
press frame; f, stand; g, inlet; h, outlet; i, filter cloths). (Ulrich, 1967.)
Conclusions 505
Filter plate ---i~~~m

Filter
cloth
Oil inlet
Oil outlet - ~~~~;l
Chamber press
Filter plate
Oil inlet _ ~~~~~~
Filter cloth
AAA
_--t!1'i
Oil outlet - l:~t~t~

Distance frame
Frame press
Oil inlet - .g:~g~f- Oil inlet -

~
I"- '/
Oil outlet Oil outlet -
Chamber layer filter Frame layer filter
Figure 12.12 Filter elements of a chamber press and a frame press (Ulrich, 1967).
12.10 CONCLUSIONS
Groundnuts, for many years to come, will continue to occupy the status of
both a nutritious food and a profitable cash crop. At present, groundnut
crops are grown mainly for their kernels and the oil and the meal derived
from them. The full potential of the crop has yet to be utilized. There exists
great scope for processing the shells for economically useful purposes such
as in the manufacture of activated charcoal, biogas, alcohol, extender
resins, cork substitute and hardboards. The manufacture of adhesive glues,
fire-extinguishing liquid and water-resistant powder from groundnut press
cake has not yet been commercially exploited.
Against the background of the campaign for 'natural' and 'healthy'
foods, partially defatted groundnut foods are becoming popular and will
assume greater importance in the years ahead. Groundnut flours may
contain 55-60% protein, minerals and vitamins and some carbohydrates.
However, they are much lower in their content of the gas-forming oligosac-
charides than are, for instance, soybean and cottonseed defatted flours.
Despite their relatively low nutritional value because of their low levels of
lysine, threonine and methionine, groundnut proteins have unique func-
tional properties such as low solution viscosity at relatively high concen-
trations (5-10%), good compatibility with bread dough systems, white
colour and bland flavour. In view of the above, opportunities exist for the
food industry (using extrusion texturization) to manufacture defatted
groundnut flours, groundnut protein isolates and concentrates as well as a
wide range of food products which might include vitamin-fortified infant
food, precooked dehydrated foods, groundnut bread, groundnut cheese
and groundnut milk through the extraction process. In particular, textur-
ized ground nut protein can provide an excellent yet economical substitute
for expensive animal protein needed to meet the short-term and long-term
requirements of nutritional improvement. The short-term nutritional
improvement target for Africa, for example, requires an increase in the
consumption of protein of high biological value from 11 g to 18 g per capita
per day, while the long-term target calls for an increase to 25 g per capita
per day.
The future pattern of industrial use of the groundnut crop will generally
differ from the more industrialized to the less industrialized groundnut-
producing countries. Most of the less industrialized countries have central-
ized large-scale processing facilities where groundnut oil and oil cake are
produced from the kernels. These are invariably located in urban areas, far
from the rural growing areas. In many instances the countries do not
produce enough oil to satisfy their domestic markets. In such circum-
stances, it is usually the rural areas where the deficits occur, because of the
difficulties and costs of transportation and distribution. The development
and introduction of technologies for groundnut processing on a small or
medium scale suitable for use in rural areas would help alleviate this
problem.
This concept, which is beginning to take firm root in a number of
countries, should be viewed as more than a technical solution to the need
to enhance the nutritional status of people in rural areas. It should also be
seen as a promoter of the overall development of rural areas - offering the
prospect of creating rural employment, adding value to agricultural pro-
duction, developing local engineering expertise and providing skills necess-
ary for rural agro-industrial development. With income growth,
urbanization and the need to satisfy the nutritional requirements of both
human and animal populations, more effort is expected to be put in
augmenting production as well as increasing the efficiency of post-
production activities such as threshing, shelling, storage and expelling.
Conclusions 507
In the more industrialized countries, where relevant resources are avail-
able, the emphasis is likely to be placed more on the use of biotechnology
for enhanced utilization of the groundnut crop.
The advent of biotechnology brings with it challenges and opportunities
for the groundnut crop. Biotechnology will have an impact on the future
pattern of industrial utilization of groundnut on at least three different
stages:
• The development of more efficient techniques and the enzymatic
modification of groundnut oil, such as fractioning, modifying and
transferring fatty acids.
• The development of plant biotechnology.
• The microbial production of oils and fatty acids.
These three factors will lead to more flexibility in the supply and use of the
groundnut and its oil. Also, as groundnut protein lacks essential amino
acids like lysine and methionine, biotechnological research is likely to be
carried out in order to improve the content of these amino acids. This again
will expand the range of products which can be made from the crop.
In general biotechnological research on improvement of groundnut
varieties will seek to improve yield and adaptation, enhance oil quantity,
modify oil quality, and improve quantitatively and qualitatively changes in
meal protein. However, the extent of utilization of future biotechnological
research results will depend not only upon the biological characteristics of
the crop but also on the socio-economic structure of production. The more
established, large-scale groundnut growers in countries like the USA,
Brazil, Argentina, India and China are expected to be more receptive to
new high-technology inputs for production and processing. Small, less
established growers, on the other hand, due to their very limited resources
and inadequate facilities, are not likely to be able to exploit quickly and
fully the benefits that biotechnology might bring.
With biotechnology research on other oilseed crops gaining momentum,
it is uncertain what the demand for specific oil/protein crops and their
products will be in future. But what is almost certain is that improvements
in curing, storage, breeding and aflatoxin elimination will lead, in a
measurable degree, to the use of more efficient known processing techno-
logies, and also 'new' technologies such as enzymatic processing, extrusion
texturization and biofermentation - and hence to additional and new
groundnut products. Globally, it is suggested that consumption and indus-
trial utilization of the groundnut crop will increase in future, partly due to
the expected increase in consumption of ready-to-eat groundnut products
in the former Eastern Bloc countries, including the former Soviet
republics.
Groundnuts still have much to contribute to the food and industrial
requirements of the world whether the need be for calories, proteins, fats,
vitamins or cellulose. The science of biotechnology, which itself is experi-
encing a revolution, has the potential to precipitate a revolution in the
production of the groundnut crop and in its industrial and domestic
utilization.
REFERENCES
Adams, F. and Hartzog, D. (1980) Peanut Science, 7, 120-123

Asiedu, J.J. (1987) Some selected engineering unit operations, in Agricultural Processing
Manual, University of Malawi.
Coulson, J.M., Richardson, J.F., Backhurst, J.R. and Harker, J.H. (1978) Chemical
Engineering. Vol. 2: Unit operations, 3rd ed, Pergamon Press, Oxford.
DEBS (1984) Manufacturer Confectioner, 63 (4), 53.
Duke, J.A. (1981) Handbook of Legumes of World Economic Importance, Plenum Press,
New York.
Gibbons, R.W. (1980) The ICRISAT Groundnut Program, in Proceedings of the Workshop
on Groundnut, ICRISAT, Patancheru, India, pp. 12-16.
Gutcho, M. (1973) Prepared Snack Foods, Noyes Data Co., Park Ridge, New Jersey.
Janssen, F. (1978) Janssen's Tips, p. 57.
Johnson, A.H. and Peterson, M.S. (1974) Encyclopedia of Food Technology, AVI Publishing
Company, Inc., Westport, Connecticut, pp. 682-684.
Khon Kaen University (1987) Final Report: Groundnut Sheller/Stripper Project, submitted to
the International Development Research Centre (IDRC), Canada.
Koo, E.C. (1942) Expression of vegetable oil: A general equation of oil expression. Journal
of Chemical Engineering, China, 34, 342-345.
Labuza, T.P. (1982) Shelf-life Dating of Foods, Food and Nutrition Press, Westport,
Connecticut.
Matz, S.A. (1976) Snack Food Technology, AVI Publishing Co., Westport, Connecticut.
McGraw-Hill Encyclopedia of Science and Technology (1982) 5th edn, McGraw-Hill, Inc.,
USA.
Minifie, B.W. (1964) Chocolate, Cocoa and Confectionery, Leonard Hill, London.
Ulrich, H. (1967) Mechanische Verfahrenstechnik, Springer Verlag, Berlin.
Weiss, T.J. (1983) Food Oils and their Uses, AVI Publishing Co., Westport, Connecticut.
Wilson, R.J. (1975) The market for edible groundnuts, Tropical Products Institute Bulletin
G96, TPI, London.
Woodroof, J.G. (1983) Peanuts: Production, Processing, Products, 3rd edn, AVI Publishing
Co., Westport, Connecticut.
Woodroof. J.G., Cecil, S.R. and Thompson, H.H. (1945) The effects of moisture on peanuts
and peanut products, Bulletin 238, Georgia Agricultural Experiment Station, Experiment,
Georgia, USA.
Zamula, E. (1985) Federal Drug and Administration Consumer, February 1985, pp. 24-20.
CHAPTER 13
Mycotoxins in groundnuts, with

special reference to aflatoxin
].1. Keenan and G.P. Savage
13.1 INTRODUCTION
In 1961 reports started to appear of heavy losses in turkey poults and

ducklings during the previous year. In the United Kingdom, 100000 birds
were reported to have died in approximately 500 outbreaks of what was
named Turkey X disease. Infected birds showed characteristic histological
lesions in the liver, which were initially thought to be related to seneciosis
in fowls (Siller and Ostler, 1961). The common factor in the UK outbreaks
was a diet containing 10% of certain consignments of Brazilian groundnut
meal (Blount, 1961). During the same year a similar condition was
reported in ducklings in Kenya and was traced to locally grown expeller-
processed groundnut meal (Asplin and Carnaghan, 1961). Indian ground-
nut meal, making up 6% of a diet, was linked to more deaths in the
following year (Carnaghan and Sargeant, 1961). Also in 1961, a toxic
factor was extracted with chloroform from the Brazilian meal (Allcroft et
al., 1961) and was shown to be extremely toxic for young ducklings,
producing lesions histologically identical to those in Turkey X disease
(Asplin and Carnaghan, 1961). It was also shown to be free from plant
alkaloids (Sargeant et al., 1961a). Toxicity testing of the Indian meal
(Carnaghan and Sargeant, 1961) produced liver lesions identical to those
produced by the Brazilian and East African toxic meals, though the
toxicity of these samples was considerably less.
This prompted Sargeant et at. (1961b) to examine groundnuts imported
from other areas. The toxin was found in samples from Nigeria, French
West Africa, Uganda, Tanganyika, Ghana and The Gambia, the level of
toxicity falling between the very toxic Brazilian and the moderately toxic
Indian groundnuts. That the signs and symptoms were clinically and histo-
logically indistinguishable from ragwort poisoning raised the question of
incorrect diagnosis of poisoning incidents prior to 1961. This was answered
510 Mycotoxins in groundnuts - aflatoxin
when it was shown that certain batches of Diet 18 (Paget, 1954), containing
15% groundnut meal, were associated with a non-infectious disease
encountered in laboratory animals from 1951 onwards. Trials carried out
by Paterson et al. (1962) with 15% toxic groundnut meal in an experimen-
tal diet (Rosetti Diet 18) produced results identical to those observed by
Paget (1954).
Extensive studies were undertaken to identify the toxin and its source. A
sample of highly toxic kernels, observed to be heavily contaminated with
fungi, was cultured. Of the eight fungal isolates grown, only one produced
an extractable toxin. This was subsequently shown to produce the typical
liver lesions. The fungus was later identified as Aspergillus flavus Link ex
Fries (Sargeant et al., 1961c). It is now known that the groundnut seed
(Arachis hypogaea L.), which develops in the soil, is particularly suscep-
tible to invasion by fungi during its development. A. flavus and A. parasiti-
cus Speare are two closely related species of ubiquitous soil fungi found in
the tropics and subtropics. Certain environmental conditions predispose
these two fungi to proliferate in the soil, with subsequent invasion of the
ground nut (Pettit, 1986; Cole, 1989). Some strains have the ability to
produce toxic metabolites, known as aflatoxins B1, B2, G1 and G2 (AFB1,
AFB2, AFG1 and AFG2). They also have carcinogenic, mutagenic and
teratogenic properties.
13.2 NATURAL OCCURRENCE OF AFLATOXINS
The four different aflatoxins are produced in widely varying amounts and
proportions depending on the genetic capabilities of the fungus, the plant
variety and the environmental conditions (Goldblatt, 1971; Jarvis, 1971;
Schindler et al., 1967; DiProssimo, 1976). The toxigenic strains generally
produce only two or three aflatoxins under any given set of conditions.
AFB1 is always produced; AFG1 is produced by the oxidation of AFBl.
AFB2 and AFG2 are only ever produced in small amounts. They were
originally thought to be produced from their respective precursors, AFBI
and AFGl. Yabe etal. (1988) showed AFB1 and AFG1 to be formed from
different precursors to those required for AFB2 and AFG2 production.
Moisture, either as kernel moisture content (KMC) or relative humidity,
has been shown to be the most important environmental parameter in the
production of aflatoxin, followed closely by temperature. Different opti-
mum temperatures have been shown for both the growth of the fungus and
aflatoxin production, including changes in the ratio of AFB 1 to AFG 1
(Jarvis, 1971). Trace elements, particularly zinc, are also required for
growth and sporulation of fungus, as well as for toxin production (Tupule,
1969). A toxigenic strain utilizes lipids in the biosynthesis of aflatoxin by
methyltransferase and oxidoreductase enzymes (Yabe et al., 1988). A zinc
Pre harvest contamination 511
or copper deficiency thus causes an accumulation of lipid in toxigenic
strains, rather than the synthesis of aflatoxin (Denning, 1987).
The competitive growth of fungi is reported to result in lower toxicities.
Several studies (Joffe, 1969; Jarvis, 1971; Tsubouchi et al., 1983) have
shown non-aflatoxin-producing A. flavus, A. niger, Rhizopus spp. and
strains of Nocardia to degrade aflatoxin. Nakazato et al. (1990) have now
shown that non-aflatoxin-producing fungi also have the ability to reconvert
aflatoxicol to AFBl.
13.3 PREHARVEST CONTAMINATION
Preharvest contamination is a major economic problem for the groundnut

industry. This was supported by a study of various food crops by Schroeder
and Boller (1973) which showed groundnuts to be the most susceptible and
most heavily contaminated of those studied. There are two types of
preharvest aflatoxin contamination in the crop. Groundnuts mechanically
or biologically damaged in the soil are predisposed to invasion by fungi.
The saprophytic fungus will live predominantly on dead or dying tissue,
therefore it may infect the maturing kernel if the pod is damaged while still
in the ground, especially when the growth rate of the plant is in decline
during the later growing phase. Hot, dry soil conditions predispose to
insect damage by termites which are known carriers of A. flavus spores.
Alternate periods of rain and drought can lead to burst pods. High levels of
aflatoxin are characteristically found in damaged kernels.
Another type of preharvest invasion can occur with no obvious kernel
damage. An extensive series of studies has provided considerable insight
into the cause of this (Blankenship et al., 1984; Cole et al., 1985; Dorner et
al., 1989; Hill et al., 1983; Sanders et al., 1985a).
The groundnut plant is unusual: it flowers above ground and, following
fertilization, the peg is pushed into the soil and the fruit develops beneath
the surface of the soil. Cole et al. (1986a) used colour mutants of both
aflatoxin-producing strains, together with aflatoxin analysis of kernels, to
prove that groundnuts are infected in the soil and not via flowers or aerial
pegs. The study strongly indicated that the most significant route of
invasion occurs after the groundnut peg penetrates the surface of the soil,
some time during fruit development. The toxigenic fungi then remain
inactive or dormant and under normal environmental conditions there is
no significant aflatoxin contamination. This is because actively growing
plants have a natural protective mechanism against disease. They are able
to produce and accumulate phytoalexins, which are antimicrobial and
antifungal compounds of low molecular weight (Strange, 1984). Three
different phytoalexins have been found in the kernels and leaves of
groundnuts. These have been named arachidins I, II and III. Water activity
(Aw) of more than 0.97 in the kernel allows sufficient phytoalexin
production to inhibit A. flavus growth. (Aw is defined as a measure of the
available water in a food system, derived from the relative humidity of the
atmosphere with which the food is in equilibrium.)
As the plant metabolism slows down during the maturation process,
phytoalexin production falls. Below 0.95 Aw the plants are unable to
produce phytoalexins, regardless of the temperature or maturity of the
kernel, but fungal growth and aflatoxin production can occur down to 0.85
Aw. Cole (1989) describes this as a 'window of susceptibility' which leaves
the groundnut kernels, especially the immature forms, without their pri-
mary resistance mechanism. The kernels seem more susceptible to afla-
toxin contamination when the Aw is between 0.95 and 0.90. Below an Aw
of 0.90, growth becomes restricted by reduced availability of water.
Dorner et al. (1989) also showed that kernel moisture loss was accelerated
at 29°C, compared with 25 °C, and that this higher soil temperature
promoted fungal growth and aflatoxin production once phytoalexin-
producing capacity was lost. In support of this, Diener and Davis (1968)
reported that old, overmature or moribund ground nut tissues have con-
siderably less KMC and increased susceptibility to A. flavus.
Preharvest aflatoxin contamination occurs when there is extreme and
prolonged drought stress during the last 4-6 weeks of the growing season.
Bushnell's study in Rhodesia (1965) showed that in a drier year there was a
high level of preharvest contamination with toxigenic strains of A. flavus,
even with rapid drying of nuts. Groundnuts subjected to these conditions
during the last 3-50 days were highly contaminated although visibly unda-
maged; a stress period of less than 20 days was insufficient to cause high
levels of contamination (Cole et al., 1986b). Drought stress causes the
groundnut canopy to recede, allowing the soil temperature in the geocar-
posphere to increase. (The geocarposphere is defined as the fruiting zone
of the groundnut plant, found 5 cm below the soil surface.)
Hot, dry soil conditions can encourage a rapid A. flavus build-up,
possibly by eliminating microbial competitors. Moisture is lost from the
kernels, the natural maturation process is accelerated and eventually the
phytoalexin-producing capacity is lost (Dorner et al., 1989). The soil
temperature is also critical for aflatoxin production (Hill et al., 1983;
Blankenship et al., 1984; Cole et al., 1985; Cole et at., 1986b). Under
drought conditions, a minimum mean geocarposphere temperature of
26°C is required for some contamination of sound kernels; lowering the
temperature to 25.7°C results in no aflatoxin production. A mean geo-
carposphere temperature of approximately 26.3-30.5°C is necessary for
preharvest contamination (optimum of 29.6-30.5 0C). A mean tempera-
ture of 31.3 °C is too high.
Adequate irrigation effectively prevents preharvest aflatoxin formation,
even when soil temperatures are optimal for aflatoxin production and
approximately 50% of the kernels are invaded by A. flavus (Hill et al.,
1983; Sanders et at., 1985a). Also, A. niger grows vigorously in hot, moist
Post-harvest contamination 513
conditions. This fungus is a known antagonist to A. fiavus, with the ability
to degrade aflatoxin.
Under stress conditions immature kernels become contaminated more
easily and quickly than the larger, mature kernels that have passed beyond
a certain developmental stage. It has been postulated that the capacity to
produce phytoalexins at a lower Aw would be a good criterion for the
selection of genotypes for aflatoxin resistance. However, under normal
conditions the phytoalexin-producing capabilities of the mature kernels
have been shown to be lower than those of immature kernels. This suggests
that they possess some additional form of resistance, a theory which is
supported by their resistance to preharvest contamination, even under
conditions of severe and prolonged drought.
Groundnut cultivars vary in their resistance to invasion by A. fiavus
(Strange, 1984; Kannaiyan et al., 1989; Siwela and Caley, 1989).
Genotypes classed as resistant in the laboratory have become extensively
colonized by A. fiavus and contaminated by aflatoxins under stress con-
ditions in the field (Sanders et al., 1985b). Some genotypes have now been
found that only support very low levels of aflatoxin production when
invaded by A. fiavus but this is dependent on the testa being entire and
undamaged (Subrahmanyam et al., 1980; Subrahmanyam, 1990; Manzo
and Misari, 1987). All developing shells are susceptible to fungal invasion,
but the degree of penetration through the pod cavity has been shown to
vary between cultivars. Kernels possessing a thick waxy seed coat and
small, covered hila are more resistant to invasion (Pettit, 1986; Pettit et al.,
1989). The presence of tannin-like compounds in the seed coat and inhibi-
tory compounds in the cotyledon also have an inhibitory effect on A. fiavus
growth and aflatoxin production. It is thought that these features, together
with the concentrations of phytoalexins and other compounds, are respon-
sible for the variation in resistance found between cultivars (Azaiaeh et al.,
1990).
13.4 POST-HARVEST CONTAMINATION
While groundnut hulls may be invaded by A. fiavus during growth in the

soil, there is also potential for contamination to occur during the drying,
transport and storage stages. Once again moisture is the single most
important parameter, followed closely by temperature (Austwick and
Ayerst, 1963; Llewellyn et al., 1983; Llewellyn et al., 1988).
At maturity the whole plant is uprooted and left in a windrow on the soil
surface to dry naturally. (Pods left in the ground beyond maturity show a
gradual increase in fungal invasion of the seeds.) Depending on conditions,
it can take 2-3 days for the moisture content of the kernels to be reduced
from 40-50% to 20-25%. The more quickly natural drying is achieved after
lifting from the soil, the less aflatoxin contamination is observed. However,
514 Mycotoxins in ground nuts - aflatoxin
care must be taken that the nuts do not dry too rapidly. This can weaken
the seed testa and testa damage decreases resistance to fungal penetration
(Manzo and Misari, 1987).
Once the pods are field-dried, they are stripped from the plants. They
should then be either sun-dried or artificially dried to <9% moisture prior
to storage. The importance of this has been shown by the much higher
incidence of kernel contamination in stored groundnuts harvested during
the wet season and insufficiently dried (Tupule, 1969; Habish et al., 1971).
Another factor that can lead to increased levels of contamination is that
groundnuts are often sold at this stage - higher moisture levels mean more
weight and increased profits for the grower.
Once dried to <9% KMC, groundnuts can be stored for long periods
with little loss of quality if kept under controlled storage conditions, i.e.
adequately cooled and well-ventilated (Baur, 1975; Aibara et al., 1985).
These conditions will also prevent the spread of any preharvest contami-
nation. Groundnuts stored in the shell are less likely to become contami-
nated during storage, due to the presence of the seed testa (Xiao, 1989). In
a 2-year study of naturally contaminated groundnut materials, Bauer
(1975) showed no statistically significant changes in aflatoxin levels with
storage.
The use of modified atmospheres at ambient temperatures for bulk
storage has been investigated with some success (Sanders et al., 1968).
While it may offer a way to control aflatoxin accumulation by inhibiting the
growth of toxigenic fungi, another study by Wilson et at. (1985) showed
changes in the microflora of the stored nuts, depending upon the KMC and
the modified atmosphere used. They also showed that something simple
like diurnal temperature changes can cause moisture migration. This
resulted in an increase from the 6-7% KMC in the original bulk sample to
one of 11 % in the top layer of ground nuts stored under 60% CO 2 . This rise
in moisture in the top layer resulted in a layer of visibly mouldy
groundnuts.
13.5 PRESENCE IN FOODSTUFFS
In the USA, Canada, New Zealand and many European countries, regu-
lations are applied to aflatoxin levels in imported groundnuts (Van
Egmond, 1989). Most have declared a tolerance level of 5 f.Lg/kg for
aflatoxin Bl and 5-20 f.Lg/kg for total aflatoxin in foods. A study in
Czechoslovakia by Fukal et al. (1987) showed only 1.9% of the samples
tested for aflatoxin gave a value above 5 f.Lg/kg. No real difference between
raw and roasted groundnuts was seen in this study. A survey in Norway by
Yndestad and Underdal (1975) showed that 17.5% of groundnuts tested
contained between 2 and 400 f.Lg/kg aflatoxin but most were at the lower
end of the range. A 6-year study in Canada carried out by Gelda and Luyt
Presence in foodstuffs 515
(1977) showed that 90.1 % of groundnuts and groundnut products had total
levels of aflatoxin <5 j.Lg/kg. In New Zealand, Stanton (1977) found afla-
toxin Bllevels ranged from 5 to 6 j.Lg/kg in two of 16 samples of imported
groundnuts. However, a further 57 outwardly clean nuts from one sample
developed A. fiavus and A. niger when incubated in agar, which demon-
strated the potential for toxin contamination during incorrect storage. The
screening of groundnuts certified aflatoxin-free from the country of origin
and imported in bulk into Trinidad showed aflatoxin levels to be <15 j.Lg/kg
(Chang-Yen and Felmine, 1987).
A study by Lim and Yeap (1966) showed one third of groundnut samples
imported into Malaysia to be contaminated with aflatoxin G 1, with no
aflatoxin Bl being detected. This raises an interesting observation. If
interpretation of the presence of aflatoxin is based solely on AFBllevels,
these samples would have been assessed as being aflatoxin free. In terms
of oral toxicity this could be significant; the oral toxicity of AFBI
(0.35 mg/kg) is only 2.9 times higher than AFG 1 (1.0 mg/kg). In contrast,
an interesting study by Strzelecki et al. (1990) assessed the occurrence of
aflatoxins in groundnut meal imported into Poland from a number of
groundnut-producing countries (mainly India and Brazil) over an 8-year
period. They found the mean contamination by AFB2, G 1 and G2 to be
9.3% of the four main aflatoxins and their toxicity to be equal to 2% of
AFB1. These experiments suggest that measuring the AFBI levels in
samples would be just as reliable, as well as being more rapid, easier and
cheaper than measuring the full range of aflatoxins.
In those countries where groundnuts and groundnut products arc locally
grown and produced, and import regulations do not apply, the biological
risk of exposure to aflatoxins is much higher owing to potentially much
higher levels of contamination. Varsavsky and Sommer (1977) reported
AFB 1 contamination in 13.1 % of samples grown and assayed over a two
year period in Argentina, with highest levels reported in the 100-1000 j.Lg/
kg range. Keen and Martin (1971) compared samples of stored groundnuts
in Swaziland and found 26% of those from rural areas to contain afla-
toxins, whereas those stored in agricultural stations were aflatoxin free
despite the presence of A. fiavus in 7.6% of samples. They attributed
the large difference in contamination to differences in storage containers
and methods of storage. Different storage practices between Fiji and
Tonga were reflected when 50% of Fijian samples were shown to be
contaminated, compared with 9% of those from Tonga (Lovelace and
Aalbersberg, 1989).
Groundnut products such as cake, meal and pellets, produced by expel-
ling oil from groundnut kernels, can be consumed locally and used in the
preparation of animal and poultry feeds. They can also contain high levels
of aflatoxin (Abalaka and Elegbede, 1982; Akano and Atanda, 1990; Lim
and Yeap, 1966; Natarajan et al., 1975a; Strzelecki and Cader-Strzelecka,
1988). Once these toxins enter the food chain, there is the potential for
secondary transmission. Eggs do not appear to contain detectable levels of
aflatoxin, even when the pullets are fed contaminated meal (Allcroft and
Carnaghan, 1963). Coker (1979) showed laying hens to excrete 70% of a
dose of AFB1, which means that <30% is passed into the eggs. Platonow
and Beauregard (1965) failed to show any significant changes in the tissues
of ferrets fed meat from chickens raised on diet containing aflatoxin. Liver
from an infected cow showed no toxicity (Alleroft and Carnaghan, 1962)
but AFM1 has been found in the liver and kidney of sheep (Reed and
Kasali, 1989). AFM1, a hydroxylated metabolite of AFB 1, has also been
found consistently in the milk of lactating animals exposed to dietary AFB1
(de longh et at., 1964; Fehr et at., 1968; Ferrando et at., 1984). This
metabolite appears 12-14 hours after ingestion of AFB1 and remains
constant throughout the feeding period, disappearing 3-4 days after the
removal of the contaminated food (Allcroft and Roberts, 1968). AFM1 has
almost identical toxicity to AFB1 but is 10 times less carcinogenic (Cullen
et at., 1987; Groopman et at., 1988; Strzelecki et at., 1990). It is recognized
as a potential health hazard and permitted levels are subject to legislation
in many countries. In the USA, the Food and Drug Administration has set
a practical action guideline of 0.5 Jl.g/kg aflatoxin M1 for fluid milk. The
major control is achieved through the regulation of AFB1 content of
animal feed.
Another commodity being utilized increasingly in animal feed is ground-
nut hulls. Sanders et at. (1984) showed that inoculated hulls can support the
growth of the fungus but not the production of aflatoxin. They also noted
that, when contamination was found in hulls from groundnuts, they were
more likely to have been machine-hulled. Hulls from the same samples
obtained by hand-shelling contained no detectable aflatoxin. This suggests
that damaged kernels or small groundnuts were being mixed with the hulls
in the machine-shelling process. Even so, they concluded that the risk
associated with the use of groundnut hulls in animal feed would be rela-
tively low.
A number of studies have been carried out on the occurrence and effect
of aflatoxins in cooking oils. In many countries unrefined ground nut oil is
consumed almost on a daily basis. For example, Fong et al. (1980) reported
that 90% of Chinese households in Hong Kong repeatedly reuse groundnut
oil for daily cooking. Studies have also shown that crude oil obtained by
hydraulic pressing contains a much higher aflatoxin content than that
obtained by solvent extraction (Parker and Melnick, 1966). Correct re-
fining of ground nut oil should give a product free of aflatoxin, regardless of
the quality of groundnuts used. The finding of aflatoxin B11evels as high as
9 Jl.g/kg in refined groundnut oil in Nigeria (Abalaka and Elegbede, 1982)
therefore gives cause for concern.
Fong's study (1980) showed aflatoxin concentrations ranging from 95 to
1055 Jl.g/kg in 10 samples of oil-grade groundnuts tested (obtained from
local markets in Hong Kong and presumably the small and immature
The chemistry and metabolism of aflatoxin 517
seeds); three oil samples had levels ranging from 98 to 150 J.1g/kg. This
correlates with a study by Dwarakanath et al. (1969) in India. They assayed
the aflatoxin content in unrefined oils prepared from freshly harvested nuts
and from those stored for 6 months and found similar levels (100 J.1g/kg and
140 J.1g/kg respectively). Another Indian study by Sarnaik et al. (1988)
showed the presence of toxigenic A. fiavus strains in five of 26 oil samples
tested. Included in this were seven samples of refined oil which showed no
contamination by A. fiavus, toxigenic or otherwise. Unfortunately this
group did not report aflatoxin levels in the contaminated samples; how-
ever, they did demonstrate that the toxin must be present in the oil seed,
by the inability of A. fiavus group fungi to produce aflatoxin in samples of
sterile oil. This indicates that the level of aflatoxin in the oil samples is
directly related to the level of contamination in the groundnuts used to
produce the oil.
The cumulative effects of the daily ingestion of small amounts of afla-
toxin on the organs of the body is not known, but the incidence of
hepatocarcinoma in humans shows a trend in parallel with aflatoxin B1
consumption in certain areas of the world. Fong et al. ( 1980) demonstrated
mutagenicity in unrefined oil, related mainly to AFB1 content.
Dwarakanath et al. (1969) found that the weight of oil absorbed by food
fried in contaminated oil increased with the weight of the food; corre-
spondingly the quantity of toxin in the food increased. However, studies
also showed that repeated cooking with the oil decreased the aflatoxin
content; e.g. heating contaminated oil at 120°C for 10 minutes resulted in a
50% destruction of the aflatoxin content. Maximum destruction of toxin
occurred in 20 minutes at 150 0c. A higher heating temperature was
needed, rather than heating longer at this temperature, to further reduce
levels.
13.6 THE CHEMISTRY AND METABOLISM OF AFLATOXIN
The aflatoxins are a family of difuranocoumarins (Figure 13.1) produced as

secondary metabolites by strains of A. fiavus and A. parasiticus. AFB1 and
B2 were named because of their strong, blue fluorescence under ultraviolet
light; whereas G1 and G2 fluoresce greenish-yellow. The B group are
characterized by the fusion of a cyclopentenone ring to the lactone ring of
the coumarin structure. The G group contain an additional lactone ring
(Groopman et al., 1988).
AFB1 (the most toxic and carcinogenic) and AFG1 both possess an
unsaturated bond at the 8,9 position on the terminal furan ring. AFB2 and
AFG2 lack this double bond. AFG1 is about 18% as active as AFB1 in a
DNA-binding assay (Gurtoo et al. 1978), whereas AFB2 and AFG2 are
essentially biologically inactive unless first metabolically oxidized to AFB1
and AFG1 in vivo (Groopman et al., 1988).
1~ 1~
aflatoxin 81
CQCXOCH3 HO~OCH3
aflatoxin B2
aflatoxin 82a
1~
ccCL~
o 0 3
aflatoxi n G 1
aflatoxin G2
aflatoxin 82a
o 0
9H (1J
c:xJCXoc:
aflatoxin M2
aflatoxicol
aflatoxin PI
aflatoxin 01
Figure 13.1 Structure of aftatoxins.
Aflatoxins are oxidatively metabolized by the microsomal mixed-

function oxygenase system mainly localized on the endoplasmic reticulum
of liver cells. This results in the formation of various hydroxylated deriva-
tives (Figure 13.2), as well as an unstable, highly reactive epoxide metab-
olite, AFBI-8,9 epoxide. It is the epoxide metabolite (which can react
covalently with the nucleophilic centres in cellular macromolecules such as
DNA, RNA and protein) that is directly related to the susceptibility of a
species to AFB I-induced hepatocarcinogenesis. A number of comparative
studies have been made on the in vitro metabolism of AFBI (Portman et
at., 1968; O'Brien et at., 1983; Ramsdell and Eaton, 1990; Lipsky et at.,
1990). A microsomal enzyme, cytochrome P-450, has been shown to be the
catalyst (Shimada and Guengerich, 1989). Roebuck and Wogan (1977)
postulated that different cellular concentrations of the same microsomal
enzyme could exist between species and individuals of a given species, to
account for the differing effects of the toxin. Since then multiple species of
P-450 have been shown to occur in hepatic mitochondria, as well as
considerable (lO-fold) inter-individual variation (Shayiq and Avadhani,
1989).
Mouse microsomes have a higher activation ratio (defined as the rate of
epoxide formation divided by the sum of the rates of the oxidative AFBI
metabolites formed) than rat microsomes. However, they are resistant to
ACTIVATION DETOXIFICATION
o 0
aflatoxin Q 1 potential exists

cytochrome P450
1'<::: 1 aflatoxin Ml for activation to
aflatoxin PI reactive epioxide
o 0 .& OCH 3
aflatoxicol
aflatoxin Bl
cc60 Icytochrome P450
+ 0 0 o 0
~
glutathione-S-transferase + GSH HO" ((
o
OCH 3 G-S' ~O' ~O' '-:Y ~OCH3
aflatoxin BI -8,9 epoxide aflatoxin B1-GSH
! 0 o 0
HO, r.' HO, /

I i Ir
o
HN~N HO' ~O' ~O' '-:Y ~OCH3
AFB1-dihydrodiol 8,9-dihydro-8,9-dihydroxy-aflatoxin B·
H2N A NJl N) o 0
o o / DNA
~ HO, ___
i ..
HO, ", o
o I II
J1y,W ~O
HN~N HN~.. )l 'CHO
H2 N AN t-;JH
H2N
A ..N Jl4
N DNA
AFB1-N 7-Gua 8,9-dihydro-8-(N 7-guanyl)-9-hydroxyaflatoxin Bl AFB1-FAPyr aflatoxin Brformamido-pyrimidine
Figure 13.2 Metabolic transformations of aflatoxin B I'

the carcinogenic effects of AFB 1 due to the efficient conjugation of epox-
ide with GSH (reduced glutathione) (O'Brien et ai., 1983; Ramsdell and
Eaton, 1990). The levels of glutathione S-transferase in mice are high
enough to convert 12 times more epoxide to AFB1-GSH compared with
male rats. O'Brien et ai. (1983) also found female rats to produce less
epoxide and to have higher levels of glutathione S-transferase than male
rats, which would help account for the decreased susceptibility of females
rats to primary liver cancer (PLC).
Several drugs and conditions have been shown to influence the levels
and activity of cytochrome P-4S0. It is highly responsive to changes in diet
and to inducers, e.g. phenobarbital. Rats fed a low protein diet became
more sensitive to the acute lethal effects of aflatoxin (Tupule, 1969).
Correspondingly, those fed aflatoxin, plus phenobarbital as an inducer,
responded in such a way that suppressed the carcinogenic activity of a
large portion of the ingested aflatoxin (McLean and Marshall, 1971).
While phenobarbital treatment has been shown to increase AFBI inacti-
vation by hydroxylation, this route is now thought to playa minor role.
LotIikar et ai. (1989) showed that rats fed aflatoxin plus phenobarbital as
an inducer have much higher levels of glutathione S-transferase.
Therefore, in species that convert more aflatoxin to the epoxide (e.g. rat
and mouse), the amount of hepatic glutathione S-transferase activity is
inversely related to the susceptibility of the species to AFBl-induced
hepatocarcinogenesis.
The AFBl-8,9 epoxide interacts with the 7 position of guanine in DNA
but spontaneous hydrolysis of the glycosyl bond means it is rapidly lost
from nucleic acids in vivo (Bennet et ai., 1981). Essigmann et ai. (1982)
showed acute doses of the toxin to cause reversible inhibition of DNA and
RNA synthesis. The adduct formed, AFBI-N7-GUA [8,9-dihydro-8-(N7-
guanyl)-9-hydroxyaflatoxin Bl], accounts for more than 80% of adducts
found in the liver of rats 2 hours after administration, i.e. it is the major
adduct formed in vivo and in vitro. In rats it is excreted in urine in a dose-
dependent manner. The amount excreted over 48 hours represents a
relatively fixed proportion (30-40%) of the total amount of AFBI-N7-
GUA initially formed in hepatic DNA (Bennett et ai., 1981; Autrup
et ai., 1987). Another adduct, AFBI-FAPyr [aflatoxin Bl-formamidio-
pyrimidine], has also been found (Lee et ai., 1989). This imidazole ring-
opened adduct, the persistent form of AFBI-N7-GUA, has been found in
liver tissue and may play an important role in hepatocarcinogenesis (Hsieh
et ai., 1988). AFBl-8,9 epoxide can also bind to protein. It has been shown
that approximately 2% of ingested AFBI binds covalently to lysine
residues in albumin, i.e. it binds quantitatively in relation to dose (Wild
et ai., 1990). Some epoxide is hydroxylated to AFBl-8,9-dihydrodiol and
excreted.
In humans, AFQl is the major aflatoxin metabolite formed from AFBl,
comprising 70-90% of the soluble metabolites (Roebuck and Wogan,
The chemistry and metabolism of aflatoxin 521
1977; Moss and Neal, 1985; O'Brien et al., 1983; Lipsky et al., 1990;
Ramsdell and Eaton, 1990). AF-8,9 dihydrodiol (10-30%), AFM1 and
aflatoxicol (AFL) are also detected; AFP1 is not present in measurable
amounts (Wild et al., 1990). The formation of these hydroxylated metab-
olites, rather than the reactive epoxide metabolite, means the production
of compounds with substantially lower carcinogenic potency than the
parent AFBl. However, the hydroxylated metabolites retain the un-
saturated bond at the 8,9 position of the molecule, which means that the
potential exists for further oxidation to an epoxide (Essigmann et al., 1982;
Groopman et al., 1985). Some are partially detoxified by enzymatic conju-
gation with sulphate or glucuronic acid to form water-soluble sulphate or
glucuronic esters, before being excreted in bile or urine (Groopman et al.,
1988). AFL, formed by an AFB1 reductase, can be reconverted to AFB1
by a dehydrogenase enzyme. Species with a high sensitivity to AFB1
appear to have a high ratio of AFB1 reductase activity to AFL dehydro-
genase activity (Salhab and Edwards, 1977).
Of these hydroxylated metabolites, AFM1 is of particular interest. First
discovered in cow's milk (Allcroft and Carnaghan, 1962), it has been
shown to have a similar acute toxicity to AFBl(O.32 mg/kg). Up to 3%
of ingested AFB1 is excreted in the milk of lactating animals as AFM1
(de longh et al., 1964; Fehr et al., 1968; Ferrando et al., 1984; Wild
et al., 1987). While proven to be a carcinogen in male rats and rainbow
trout, it is a weak hepatic carcinogen compared with AFB1 (Cullen et al.,
1987).
Humans are capable of producing AFB-DNA adducts but at lower
levels than rat cells. The fraction of AFB1 activated by human liver
microsomes increases with decreasing AFB1 concentration. This is due to
the high affinity for AFB1 of the cytochrome P-450 isoenzyme that forms
the -8,9 epoxide, compared with the low AFB1 affinity of the AFQ1-
forming species of enzyme. Compared with some species studied, human
liver glutathione S-transferase appears to lack the substantial capacity for
epoxide conjugation found in rats and mice (Moss and Neal, 1985) there-
fore the liver tissue may be relatively efficient at generating AFB-DNA
ad ducts when exposed to dietary levels of AFB 1 (Ramsdell and Eaton,
1990). Both rat and human hepatocytes display nucleolar segregation
proportional to the level of AFB-DNA binding. This is thought to be a
morphological manifestation of carcinogen-DNA binding (Lipsky et al.,
1990).
As with rats, human males have been shown to metabolize the aflatoxin
2 to 3 times faster than females. A 16-fold inter-individual variation in the
level of AFB-DNA adduct formation has been shown (Lipsky et al., 1990).
There is also the possibility that the pattern of aflatoxin metabolism in
compromised livers (damaged by alcohol or hepatitis B virus) may differ
from that in normal livers.
TABLE 13.1 Acute toxicity of AFB] with a single dose (LD50 mg/kg)
Wogan Newberne Roebuck Coker

(1968) and Butler and Wogan (1979)
(1969) (1977)
Duckling 0.4 0.335 0.3

Rainbow trout 0.5 0.5-1.0
Pig 0.62 0.6
Baboon 2.0
Chicken 60-16.0
Rat (male) 1.0 0.56-17.9 1.0-5.0 7.0
Rat (female) 18.0
Mouse 63.0 9.0 15.0 9.0
Rabbit 0.5
Dog 0.5 1.0
Guinea pig 1.0 1.4
13.7 AFLATOXICOSIS IN ANIMALS AND POULTRY
Aflatoxin B1 is the most toxic of the naturally occurring aflatoxins. It is

reported to be almost twice as toxic as strychnine and potassium cyanide to
rats (Strzelecki et ai., 1990). In all species studied, the liver appears to be
the target organ, resulting in toxic hepatitis, cirrhosis or primary liver
cancer (Reed and Kasali, 1989). The time taken for animals to succumb to
aflatoxin appears to be related to the total dose of the toxin, rather than the
rate of consumption.
The LDso of aflatoxin for most species of animal (Table 13.1) is within
the range 0.3 mg/kg to 10 mg/kg body weight (Wogan, 1968; Newberne
and Butler, 1969; Roebuck and Wogan, 1977; Coker, 1979; Akano and
Atanda, 1990). The observed range of LDso values emphasizes the fact
that the metabolism of aflatoxin varies quantitatively and qualitatively
between animal species, thus causing marked differences in their suscepti-
bility to the toxin. The tolerance to an acute (or subacute) dose can also
vary widely within a given species. Factors governing this can include the
age, sex, strain and condition of the animal, the level and type of aflatoxin
in their diet and the time elapsed before the LDso is measured (Coker,
1979). Also, not all animals of a similar age on identical contaminated
feeds are affected, which suggests genetic restriction (Loosmore and
Markson, 1961; Sisk et ai., 1968; Hintz et al., 1967; Murthy et al., 1975).
For all species of animal in which age has been recorded, the young have
been shown to be more susceptible (Loosmore and Harding, 1961; Allcroft
and Carnaghan, 1963; Horrocks et al., 1965). Females are less susceptible
than males (Wogan, 1968).
Subacute or chronic exposure affects growth feed efficiency and general
well-being, especially during the early growing period (Duthie et al., 1966).
Atlatoxicosis in animals and poultry 523
In the early stages it may not be associated with overt clinical symptoms. It
eventually leads to the development of a toxicity syndrome and liver
damage. Clinical recovery can occur when the contaminated diet is
removed; however, the livers of the affected animals will still show
aflatoxin-induced changes (Duthie et al., 1968).
Calves and pigs are the most susceptible domestic animals. Dairy cows
are reported to have significantly reduced milk yields which coincide with
the excretion of AFM1 in their milk (Allcroft and Carnaghan, 1963;
Coker, 1979). Ray et al. (1986) reported abortion and symptoms suggestive
of acute aflatoxicosis after pregnant cows ingested aflatoxin-contaminated
groundnuts. Pigs fed aflatoxin during gestation did not abort but did show
histological changes in their livers. In this study no biochemical effects
were detected in their offspring (Gumbmann and Williams, 1969; Ray et
at., 1986). Sheep have been reported to tolerate a heavily contaminated
diet for up to 3 years (Wogan, 1968). Some species, e.g. monkeys, calves
and rabbits, can develop cirrhosis after chronic exposure. Chickens are less
susceptible than turkey poults; ducklings are the most susceptible species
known, which makes them ideal for biological screening purposes (Asplin
and Carnaghan, 1961).
Both lipid and protein metabolism are affected by aflatoxin ingestion,
manifested by an early decrease in feed conversion. The biochemical
changes in the affected animals appear to be closely related to the degree
of hepatic injury, which is in turn related to dose administered. The
vacuolization and fatty changes which occur in the hepatocytes lead to
necrosis, which is usually localised in one part of the hepatic lobule,
depending on the species (Newberne and Butler, 1969; Syamasundara
Rao, 1970). This accumulation of fat, seen in the early stages, is thought to
be due to inhibition of fat removal from the liver (Reed and Kasali, 1989).
Bile duct proliferation is a typical pathological change; hepatic veno-
occlusive lesions are also common.
Because of its ability to modify DNA, AFB1 can cause a reduced rate of
protein synthesis, which in turn increases the protein requirements of
susceptible animals. Serum albumin decreases and gamma-globulin in-
creases (Annau et at., 1964). Therefore animals with an already reduced
protein intake would have increased susceptibility to aflatoxin, due in part
to the inability of the liver to synthesise the enzymes necessary for the
metabolism and detoxification of the aflatoxin, leading to longer exposure
and increased toxic effects (Tupule, 1969; Coffey et al., 1989). Increasing
the level of dietary protein or lipids above the normal requirement for
optimum performance has been shown to prevent depressed performance
due to aflatoxin exposure (Smith et al., 1971). Coffey et al. (1989) showed
increased dietary lysine or methionine can protect against aflatoxicosis.
This is of special interest because it is now known that aflatoxin binds to the
lysine residues of plasma albumin in a dose-related manner (Wild et al.,
1990).
Aflatoxin is also known for its immunosuppressive effect. Coker (1979)
reported that poultry fed diets containing 250-500J.Lglkg were predisposed
to attack by viruses and bacteria. This secondary effect could be particu-
larly significant in livestock already compromised by poor nutritional status
(Reed and Kasali, 1989).
The potential carcinogenicity of AFB1 is related to the proportion that
ultimately binds to DNA. This is determined by both the amount activated
in vivo and how much is subsequently detoxified (Ramsdell and Eaton,
1990; Lipsky et at., 1990). Rats, which do not show signs of acute toxicity,
are highly susceptible to aflatoxin-induced hepatomas (Lancaster et at.,
1961; Butler and Barnes, 1968). Rainbow trout and ducks are also highly
susceptible (Wogan, 1968). Liver cancer has now been described in many
species of vertebrate, although the dose and period of exposure varies
widely. It is thought that chronic exposure to small doses of aflatoxin is
more likely to produce hepatocellular carcinoma than exposure to the
same amount in larger doses over a brief period of time (Wogan and
Newberne, 1967). Even so, there are reports of rats surviving an LD50 dose
and subsequently developing hepatoma (Carnaghan, 1967; Angsubhakorn
et at., 1990). Mice appear to be resistant to both subacute toxicity and
carcinogenicity (Wogan, 1968).
A greater susceptibility of neonates to the carcinogenicity of B 1 has been
suggested (Cullen et at., 1987). Males tend to develop tumours at a faster
rate (Wogan, 1968). A difference in susceptibility between two strains of
rat has also been demonstrated (Fong and Chan, 1981; Angsubhakorn et
at., 1990).
13.8 AFLATOXIN AND HUMAN DISEASE
Within a popUlation, the character and intensity of human exposure to

aflatoxin may vary depending on factors such as age, nutritional status,
concurrent exposure to other agents (e.g. hepatitis B virus) as well as the
level and duration of exposure (Dichter, 1984). Differences between indi-
viduals in a population in AFB1-FAPyr adduct formation (the persistent
adduct) in liver tissue also suggests there is genetic restriction (Lee et at.,
1989).
The liver is the organ primarily affected by aflatoxin ingestion. The
known metabolic effects of aflatoxin ingestion include inhibition of DNA,
RNA and protein synthesis, reduction of certain enzyme activities, de-
pression of glucose metabolism, inhibition of lipid synthesis and depression
of clotting factor synthesis. Aflatoxins also tend to block steroid binding
sites and are immunosuppressive. The pathological changes include fatty
infiltration, biliary proliferation and toxic necrosis of the liver in acute
severe poisoning. There are only four outcomes to toxic damage to the
liver: recovery, cancer, cirrhosis and ongoing hepatitis (Coady, 1976). In
Aflatoxin and human disease 525
man, studies have implicated aflatoxin in the aetiology of acute hepatitis,
cirrhosis in malnourished children, kwashiorkor, Reye's syndrome and
primary liver cancer (Denning, 1987, has written an excellent review).
To date there have been only a few documented cases of acute aflatoxi-
cosis in man. These have ranged from an individual case to one outbreak in
India in which 106 people were reported to have died (Serck-Hanssen,
1970; Ngindu et al., 1982; Krishnamachari et ai., 1975). The livers of those
affected showed signs of toxic hepatitis. AFBI was found in the sera of two
out of seven patients tested for the toxin (Krishnamachari et al., 1975).
Ngindu et ai. (1982) reported AFBI (38 and 89 J.Lg/kg respectively) in two
liver samples obtained. Contaminated foodstuffs other than groundnuts
were implicated in each outbreak. Not all people exposed to the toxin were
adversely affected.
The finding of aflatoxins in the breast milk, neonatal cord blood and the
sera of pregnant women indicates that a high proportion of at-risk infants
have not only prenatal but also continuing post-natal exposure to aflatoxin
(Robinson, 1967; Lamplugh et al., 1988; Denning et al., 1990). Studies also
indicate not only transplacental transfer but also concentration of aflatoxin
by the foeto-placental unit (Lamplugh et al., 1988; Denning et ai., 1990).
Tupule (1969) showed rats and monkeys fed a protein-deficient diet were
more susceptible to aflatoxin; therefore the combined exposure of a human
foetus or infant to aflatoxin and malnutrition must severely diminish its
chances of survival.
13.8.1 Kwashiorkor
Protein energy malnutrition (PEM) is implicated in increased susceptibility
to aflatoxin by reducing the levels of cytochrome P-450 and therefore
microsomal hydroxylation of aflatoxin in the liver (McLean and Marshall,
1971). In humans PEM is divided into two main groups: marasmus and
kwashiorkor. Marasmus is due to chronic near-starvation. Kwashiorkor
has an obscure pathogenesis; it is characterized by hypoalbuminaemia,
fatty liver and immunosuppression, and is thought to be due to the
deprivation of protein in the presence of adequate calorie intake (Anon.,
1984). These metabolic and pathological consequences of kwashiorkor
closely resemble those of aflatoxin ingestion. The occurrence of kwashior-
kor also closely mirrors the geographical and seasonal prevalence of afla-
toxin contamination in foodstuffs.
Aflatoxin is not thought to be responsible for initiating kwashiorkor. It is
possible that a zinc deficiency, which becomes limiting before a protein
deficiency, precipitates the condition by interfering with protein and nu-
cleic acid synthesis. This is supported by the observation that in many cases
the liver seems to suffer little pathological damage and responds well
to zinc therapy (Golden and Golden, 1981). The ingestion of aflatoxin
could exacerbate the problem; the inability of the malnourished liver to
526 Mycotoxins in ground nuts - aflatoxin
metabolize it could lead to the more severe liver lesions occasionally seen
in children with kwashiorkor.
Breast-milk analyses have shown atlatoxins in >30% of samples, which
can account for wholly breast-fed babies showing signs of kwashiorkor
(Hendrickse, 1988). In these babies the reduced ability of the liver to
synthesize albumin could predispose them towards the kwashiorkor end of
protein energy malnutrition. Aflatoxin ingestion can be the final insult that
precipitates the child into kwashiorkor. It is also postulated that aflatoxin
causes a dose-related suppression of immune function, which would make
these children even more susceptible to infectious disease mortality (Den-
ning, 1987).
Serum measurements in children showed that aflatoxin was present more
often and in higher concentrations in children with kwashiorkor than in
those with marasmus or controls (Hendrickse et al., 1982). They also found
AFB 1 or aflatoxicol in all the autopsy liver samples they obtained from
children dying from kwashiorkor and never from those dying of other
causes. It could be argued that this is the result of greater exposure to
aflatoxin in these children but two other results have to be taken into
account. While urinary metabolites were detected most often in samples
from kwashiorkor patients, the concentrations were lower than in other
groups. Aflatoxicol, a metabolic product of AFB1 that can be converted
back to AFB1, was detected only recently in the sera of children with
kwashiorkor. This suggests a difference in metabolism in these children,
possibly linked to an inability to transport and excrete the aflatoxin
(Hendrickse, 1988). A recent study by de Vries et al. (1990) supports this
theory, and also shows that large amounts of aflatoxins can accumulate in
the body tissue of these children. Their study also suggests that the
metabolic fate of aflatoxin in children with kwashiorkor may differ from
those with marasmic kwashiorkor.
13.8.2 Cirrhosis
There is a high prevalence of childhood hepatic cirrhosis in certain parts of
India which peaks at 3 years of age (Tupule, 1969; Amlaetal., 1971). It has
been directly linked (via breast-milk studies) with Indian mothers of
cirrhotic children eating aflatoxin-contaminated foodstuffs (Robinson,
1967). Liver biopsies taken from Indian children with cirrhosis at different
stages of severity have revealed varying degrees of cell destruction,
fibrosis, inflammatory cell infiltration and regenerating nodules (Tupule,
1969). The presence of aflatoxin metabolites was also shown in the urine of
these children. Amla et al. (1971) implicated aflatoxin as a cause when they
showed that Indian children suffering from kwashiorkor, inadvertently
treated with an aflatoxin-containing diet «30 fLg/kg daily), went on to
develop cirrhosis. The hepatic lesions noted had a direct correlation with
duration of toxin ingestion, which ranged from 5 days to 4 weeks. Despite
Aflatoxin and human disease 527
this direct link with aflatoxin ingestion, studies of urine from cirrhotic
children have been inconclusive; similar levels of AFB1 have been found in
both subjects and controls (Amla et al., 1970; Yadgiri et al., 1970). This
would support the theory that childhood hepatic cirrhosis has a multifac-
torial aetiology (Coady, 1976).
13.8.3 Reye's syndrome

Reye's syndrome is a childhood disorder of previously unknown aetiology.
It is indistinguishable clinically from a condition found in Thai children,
commonest at the end of the rainy season when aflatoxin contamination of
food is highest. Both exhibit encephalopathy with fatty degeneration of the
viscera. Because of the similarity of the lesions to aflatoxin poisoning,
several studies were made on children dying from Reye's syndrome.
Aflatoxin was identified in liver samples from cases of Reye's syndrome in
New Zealand (Becroft and Webster, 1972), Czechoslovakia and the USA
(Ryan et at., 1979). In a subsequent study Rogan et at. (1985) were unable
to isolate any aflatoxin metabolites or adducts in liver biopsy samples using
more sensitive methods. Reye's syndrome is now known to occur after a
viral illness. It is possible that in some cases aflatoxin could alter the host's
response to the virus (Denning, 1987). More commonly a very strong link
has been shown between Reye's syndrome and the intake of aspirin In
children under 12 years of age (Glenn-Bott, 1987).
13.8.4 Primary liver cancer

Primary liver cancer (PLC) is a disease of striking geographic distribution
and is one of the leading causes of cancer mortality in Asia and Africa. A
number of epidemiological studies have shown what appears to be a good
correlation between the level of ingested aflatoxin and incidence of PLC
(Korobkin and Williams, 1968; Brudzynski et al., 1978; Groopman et al.,
1988), which has raised the hypothesis that exposure to dietary aflatoxin is
associated with an elevated risk of PLC. The indirect demonstration of
AFB-DNA adducts, by the identification of the AFB-Gua 1 adduct in
urine samples collected from people in different regions of the world where
there is a high incidence of PLC, adds weight to this theory (Lipsky et al.,
1990). In contrast, the incidence of PLC is low in Central and South
America, although environmental conditions match those of the high-risk
areas of Africa and Asia (Wogan, 1968).
Recent work has found aflatoxin ingestion by itself to be insufficient to
induce PLC (Stoloff, 1989; Campbell et al., 1990). This would suggest a
multifactorial aetiology, with factors such as individual susceptibility, age,
nutritional status and concurrent exposure to other agents such as alcohol
and hepatitis B virus (HBV), being important determinants in the develop-
ment of PLC. Denning (1987) postulated that selenium or some other
antioxidant deficiency could also contribute to aflatoxin-induced carcino-
genesis.
Chronic hepatitis B virus is now thought by some authors to be the major
aetiologic agent in PLC (Stoloff, 1989; Campbell et al., 1990). It is endemic
in many areas of the world in which the incidence of liver cancer is high and
PLC mortality has been shown to be positively correlated with HBsAg+
prevalence. Stoloff (1989) was even able to rule out the need for aflatoxin,
or any similar cofactor, as a necessary condition for HBV-induced PLC.
There is no disputing the fact that aflatoxin is activated in vivo by humans.
This is confirmed by the detection of the AFB1-FAPyr in liver tissue (Lee
et al., 1989) and AFB1-N7-Gua in urine (Lipsky et al., 1990). But while
aflatoxin has the potential to act as a carcinogenic initiator, consumption
must be high enough and prolonged enough for it to contribute a signifi-
cant risk (Campbell et al., 1990).
The role of aflatoxin in the aetiology of PLC is not clear. Campbell et al.
(1990) state that it is an unnecessary and insufficient cause; Stoloff (1989)
postulates that it possibly has a late-stage effect in the development of
PLC, acting in an acute (dose-dependent) manner on a liver already
compromised by HBV infection. Denning (1987) postulates that HBV may
be essential in lowering the threshold at which aflatoxin carcinogenesis
occurs. Alternatively aflatoxin may act as an immune suppressive agent,
rather than as a primary carcinogen (Lutwick, 1979). This would allow
HBV to maintain itself more easily in the liver, producing more chronic
infection. This may be critical for the initiation and development of either
PLC, cirrhosis or ongoing hepatitis. It could be that these three pathologi-
cal states may represent different responses to aflatoxin. It has also been
postulated that exposure to AFBlIAFM1 via breast milk could be relevant
in determining the integration and/or rearrangement of the HBV genome
within the cellular genome (Chandra, 1970; Wild et al., 1987).
13.9 ANALYTICAL METHODS FOR THE DETECTION OF

AFLATOXINS
Because of the potential for contamination, all groundnuts and groundnut

products should be monitored for the presence of aflatoxin throughout all
phases of their production. Originally biological methods were used to
detect the presence of aflatoxin. Sargeant et al. (1963) found that aqueous
suspensions of material extracted from toxic meals with solvents could be
administered to day-old ducklings, to measure toxicity of the sample.
Good agreement was found to exist between toxicity established by bio-
logical assay and the amount of fluorescent material determined by paper
chromatography. AFB1 was shown as a single spot, with a bright blue
fluorescence under UV light, while AFG 1 has a green fluorescence and is
less mobile.
Analytical methods for the detection of afiatoxins 529
The standard chemical method, based on thin layer chromatography
(TLC), was developed by the Association of Official Analytical Chemists
(AOAC) in 1970. High performance liquid chromatography (HPLC) is
also used for quantitative analysis. Several screening methods are now
available. Immunochemical methods, such as the Enzyme-Linked
ImmunoSorbent Assay (ELISA) and affinity column methods, are now
being developed and offered as simpler, quicker alternatives.
13.9.1 Sampling
The difficulties in obtaining a representative sample of ground nuts for
aflatoxin analysis have been established. Aflatoxin is not a natural constitu-
ent of ground nut tissue, and is present only in a proportion of kernels. In a
sample of groundnuts with a high average content of aflatoxin, only a small
fraction may be contaminated. Even these few groundnuts may have
varying levels of aflatoxin, which makes representative sampling more
difficult (Allcroft and Raymond, 1966; Cucullu et ai., 1966; Gardner et ai.,
1968). For a given aflatoxin concentration, the proportion of lots accepted
by any testing plan is dependent on the sample size, subsample size,
number of analyses, method of sample preparation and acceptable level.
The acceptable level is defined as the highest aflatoxin test result allowed
for a lot to be accepted (Whitaker and Dickens, 1989). Some countries
have a zero tolerance, which is in practice the limit of detection by the
analytical method employed (Van Egmond, 1989).
The negative binomial distribution has been used for estimating the
probabilities associated with sampling lots of shelled groundnuts for afla-
toxin analysis (Whitaker et ai., 1972). It is characterized by a continuous
decrease in the probability of finding a groundnut with increasing amount
of aflatoxin (Whitaker et ai., 1985). This model has a property which
enables distribution to be inferred for different sample sizes. However, this
property applies only if all the variability may be attributed to sampling,
i.e. variability due to subsampling and analysis may undermine the value
of the model (Brown, 1984). With the large sample sizes (three 22 kg
samples) and the high acceptance levels permitted in the USA (25 fLg/kg
total), the choice of statistical distribution model is not very important.
The tolerance accepted in Europe is much lower (1-5 fLg/kg). This, com-
bined with a small sample, can be unfavourable to the consumer, as the
chance of detecting a lot with a mean concentration greatly above the
tolerance limit will be very small (Knutti and Schlatter, 1982; Whitaker and
Dickens, 1989). The amount of sample necessary is inversely related to the
parts per billion (fLg/kg) aflatoxins (Cucullu et ai., 1966).
The particle size of the product being sampled is also a factor in
detecting the true level of the contamination present (Waltking, 1980). A
single groundnut can be severed into fewer than 50 particles to make
chunky groundnut butter. The same groundnut can be divided into more
than 300 million particles to make the creamy-style product. In the latter,
there is a high probability of there being a uniform aflatoxin concentration
(if present) which does not apply to the former.
13.9.2 Presumptive and screening methods

In the USA all of the kernels from each official grade sample (465 g) are
examined visually. If one or more kernels are found with visible A. flavus
growth, the lot is segregated for restricted oil processing, i.e. diverted from
food use (Whitaker and Dickens, 1986).
Minicolumns have also been developed as a rapid, inexpensive screening
method. They are all based on running a solvent-extract of a ground
groundnut sample through a silica gel column. Holaday (1968) developed
the original one; since then several modifications have been published
(Holaday and Lansden, 1975; Davis et al., 1980: Shotwell and Holaday,
1981). The fluorescence of the extracts under long-wave UV light (365 nm)
is then compared with that of controls. However, the test is npt specific for
aflatoxins - some fluorescence from other fungal metabolites can occur.
The Romer All-Purpose Minicolumn method has been granted official
first action for screening for aflatoxins in groundnuts (AOAC, 1984). It is
applicable for detecting ~1O /-Lg/kg in groundnuts, groundnut meals and
peanut butter and ~15/-Lg/kg total aflatoxins in mixed feeds. After extrac-
tion, the samples are treated with FeCl3 and CuC0 3 to remove interfering
substances. Lovelace and Aalbersberg (1989) showed that this method
gave a good correlation with TLC. An inexpensive modification of this step
substitutes CUS04 as the clarifying agent and still gives 100% agreement
with Romer's method (Sylos and Rodrigues-Amaya, 1989).
The Holaday-Velasco method, which has been approved official first
action for corn (AOAC, 1984), has also been recommended as a screening
method for groundnuts. It uses methanol-water extraction and benzene for
solvent partition. A collaborative trial showed that the limiting level for
reliable aflatoxin detection using this method was between 5 and 13 /-Lg/kg
(Shotwell and Holaday, 1981). A modification of this method is the
SAM-aflatoxin tube. Using toluene instead of benzene for solvent parti-
tion, this phase is then passed through the SAM (Selectively Absorbed
Mycotoxins) tube which removes other interfering compounds in a pre-
absorption layer and selectively absorbs any aflatoxin at specific bands in
the tip of the tube. The sensitivity of the assay is designed to give a yes/no
answer at designated levels of aflatoxin. It is a rapid, sensitive, economical
and chemically stable assay (Pettit et al., 1990).
13.9.3 Quantitative methods

A number of methods exist for preparing a groundnut sample for quanti-
tative analysis. The official AOAC Method I (CB method) (1984) is the
Analytical methods for the detection of aflatoxins 531
standard by which other methods are judged. Based on a chloroform-
water extraction, it gives better clean-up and higher recovery than other
methods ,(Whitaker and Dickens, 1983). This makes it ideal for use with
TLC but it is expensive and time-consuming. The AOAC Method II
(BF method) uses a methanol-water (55:45) extraction and a solvent/
groundnut ratio of 5: 1 (mIlg) for the extraction of aflatoxin from 50 g
samples of raw groundnuts (AOAC, 1984). This method is the faster and
more economical of the two.
Modifications of Method II include the water slurry (WS) method.
Whitaker et at. (1984) found the amount of aflatoxin extracted from raw
groundnuts to be a function of both the methanol concentration and
solvent/groundnut ratio. Increasing the methanol concentration to 60%
and the solvent/groundnut ratio to 10.8: 1 (ml/g) gave 12.1 % more aflatoxin
extracted. However, this increase required 2.36 times more methanol than
the original AOAC Method II (Whitaker et al., 1986). Another modifi-
cation is the Food Safety and Quality Service (FSQS) method. It has been
developed for larger sized (21.8 kg) samples and is recommended for
naturally contaminated lots where aflatoxin is unevenly produced.
A number of comparative studies have been made of the different
extraction methods (Coon et at., 1972; Mechan et at., 1985; Pluyer et at.,
1987; Van Egmonde and Wagstaffe, 1989). Minicolumn methods are also
being used as rapid clean-up methods to replace the official quantitative
methods (CB/BF), which are comparatively time-consuming and expen-
sive. The purified extracts are suitable for TLC as well as HPLC and allow
the detection of low levels of aflatoxin (Dorner and Cole, 1988). However,
Mechan et at. (1985) compared several extraction methods with the Romer
minicolumn method and found the latter to give considerably lower results.
If used, the minicolumn has the advantage in that preliminary estimation of
aflatoxin can be made which allows for dilution if necessary before quanti-
tative analysis.
(a) Thin layer chromatography

Samples for TLC are spotted onto pre coated silica gel plates. After devel-
opment, they are detected under long-wave UV (365 nm) light. A visual
comparison can be made with standards but fluorodensometric measure-
ments give a much more sensitive result. Interfering fluorescent material in
the samples, including oils, fats, sugars and protein not completely
removed from the extracts by the clean-up procedures, can cause a loss of
accuracy (Beljaars et al., 1972). This includes errors in measuring the
proportion of the solvent solution analysed for aflatoxin and in measuring
the proportion of the extracted aflatoxin placed on TLC plates (Dickens
and Whitaker, 1983). These two errors can cause underestimation of
aflatoxin content by as much as 11 %. Two-dimensional TLC procedures
have now been developed which can reduce the interference and therefore
the need for extensive clean-up procedures (Beljaars et al., 1973). In a
comparison with an ELISA estimation of aflatoxin, Chu et al. (1988)
observed that the largest error in the determination of aflatoxin by the TLC
method occurs during the silica gel clean-up and/or the TLC step itself.
(b) High performance liquid chromatography

HPLC provides a precise and sensitive method for determining aflatoxins
in all types of groundnut products. Combined with flow injection analysis it
also offers speed, simplicity and economy (Lazaro et al., 1988). Both
normal phase and reverse phase HPLC separations have been developed.
For either method samples have to be extracted and cleaned up prior to
analysis. Pons and Franz (1978) found an acidified aqueous methanol
extraction yielded consistently higher results. Samples are purified on a
small silica gel column before being eluted and quantified. Normal phase
HPLC uses a silica-gel packed flow cell for either UV detection at 360-
365 nm (AFBlIB2) or fluorometric detection (AFGlIG2). Using this
method Pons and Franz (1978) were able to resolve completely the four
aflatoxins in extracts of raw groundnuts, groundnut meal and peanut
butter. The drawback with this method is the instability of the packed cell
(Wilson, 1989).
Reverse phase HPLC separations are more widely used than normal
phase (Hurst and Toomey, 1978) but the fluorescent intensities of AFB1
and AFG1 are reduced in reverse phase solvent mixtures. Derivatization
with trifluoroacetic acid (TFA) prior to injection overcomes this by con-
verting the weakly fluorescent AFB1 and AFG1 to their respective highly
fluorescent hemiacetals AFB2a and AFG2a. Park et al. (1990) reported an
average recovery of 88.1 % and 63.8% for peanut butter and raw ground-
nuts respectively using this method. It has now been recommended as
official first action for determination of AFB1, AFB2, AFG1 and AFG2 in
peanut butter at concentrations ?o13 f.Lg total aflatoxin/kg. Another
alternative is post-column derivatization (PCD) with iodine in water to
enhance AFB 1 and AFG 1 fluorescence after reversed phase chromatogra-
phy. Dorner and Cole (1988) combined a minicolumn clean-up procedure
with this method. It proved to be sensitive, efficient and extremely rapid.
13.9.4 Immunochemical methods

AFB1 in groundnuts can also be detected by immunochemical methods
Two of these - the ELISA and the monoclonal antibody affinity column -
use specific antibodies to detect aflatoxin in different foods and feed. They
are comparatively quick, easy and potentially inexpensive. They have the
ability to detect low aflatoxin levels and the capability for routine screening
of large numbers of test samples.
The direct competitive ELISA is based on competition between an
Analytical methods for the detection of aflatoxins 533
enzyme-conjugated AFB1 and (free) aflatoxins in the test sample. They
compete for aflatoxin-specific antibodies, coated onto a solid phase,
usually a microtitre plate. A comparison by Chu et al. (1988) between the
ELISA method and each step of the TLC method showed a clean-up
treatment is not necessary in the ELISA.
When a polyclonal antibody is used to coat the ELISA plate, the result is
not confirmatory for AFBl. This is because a polyclonal antibody cross-
reacts with AFB2 (70%), AFG 1 (75%) and AFG2 « 10%) respectively
(Trucksess et al., 1989). Park et al. (1989) showed a 98% correlation
between this method and TLC results for roasted groundnuts. As such the
method has been approved as interim official first action as a screening
method to determine the presence or absence of AFB 1, AFB2 and AFG 1
in groundnut products. The detection limit is set at 20 f.Lg/kg - a positive
result for total aflatoxins at >20 f.Lg/kg or negative results at ~20 f.Lg/kg
(Park et al., 1989). Trucksess et al. (1989), investigating a modification of
this method, recommended a detection limit of >30 f.Lg/kg in raw ground-
nuts. Commercial ELISA kits are now available, providing a rapid screen-
ing method for large numbers of samples, but these kits vary in their
specificity and sensitivity (Mortimer et al., 1988; Dorner and Cole, 1989;
Patey et al., 1989).
The use of monoclonal antibodies enables the detection of very low
levels of aflatoxin. The Aflatest has been developed, based on monoclonal
antibodies adsorbed onto an affinity column, and is useful for the rapid
isolation of aflatoxins from samples of food (including milk). As the
methanol-extracted sample passes through the column, any aflatoxin in it
adheres to the monoclonal antibody. Methanol is then used to elute the
conjugated aflatoxin-antibody complex. This collects in a fluorosil tip. The
fluorescence of the sample can be compared under UV light with a set of
standards. This provides a test for total aflatoxin in minutes, using simple
equipment with a minimum of expertise (Cole and Masuka, 1989).
The affinity column also provides a rapid one-step removal of interfering
substances from diverse biological samples. The method is highly specific
and sensitive (Groopman et al., 1985) and has proved to be as effective as
the CB extraction method (Trucksess et al., 1990). The affinity column
separation can then be coupled with solution fluorometry with bromine
(SFB) to estimate total aflatoxins. Alternatively individual toxins can be
determined by reverse phase liquid chromatography with post column
derivatization with iodine (PCD). This method has now been recom-
mended as an official first action for the determination of aflatoxins in raw
groundnuts and peanut butter at total aflatoxin concentrations :? 10 ng/g
(Trucksess et al., 1991). It is interesting to note that a similar study by
Patey et al. (1991) gave much lower levels of recovery (51-67%) from
peanut butter samples compared with those reported by Trucksess et al.
(1991) for the same PCD method (81% and 83% respectively for raw
groundnuts and peanut butter).
13.9.5 In vivo detection of aflatoxin
The variability in dietary exposure to AFB1 makes it very difficult to
determine the extent of cumulative aflatoxin exposure in an individual.
Genetic susceptibility to adduct formation also makes it difficult to estab-
lish a dose-response relationship between the amount of aflatoxin ingested
and the amount of adduct found in the tissues at a given time (Lee et al.,
1989).
Aflatoxin has a short half-life in serum. It is excreted in the urine and
breast milk as a number of different metabolites which are excreted at
different rates. An ELISA method developed to detect AFB1, AFG1 and
AFQ1 in serum or plasma has proved to be a useful index of short-term
exposure, provided a blood sample is taken within 24 hours (Denning,
1987). Wilkinson et al. (1988) reported that the pretreatment of sera with a
1: 1 ratio (v/v) of methanol reduces the differing levels of interference in
different sera, caused by serum proteins. Wild et al. (1987) have developed
an ELISA for the detection of aflatoxin in human breast milk, with a
sensitivity of 2 pg AFMlIml milk. Eleven per cent of samples tested in
rural Zimbabwe gave a positive result with levels up to 50 pg/ml.
The measurement of urinary metabolites is another alternative. Samples
can be obtained easily and non-invasively. AFM1 is the major metabolite
present in urine (Campbell et al., 1970). It comprises 1-4% of ingested
AFB1 and is mostly excreted within 48 hours; therefore, it can be used as a
measure of recent aflatoxin ingestion. AFB 1 and AFP1 (in rat urine) have
also been recovered but in much smaller amounts. Measurement of
AFB1-N7-Gua in urine reflects the proportion of AFB1 activated by
exposed people (Groopman and Donahue, 1988).
The problem with measuring aflatoxin metabolites and/or nucleic
adducts in body fluids is that they are rapidly excreted and therefore reflect
a relatively short-term (few days) exposure. Another alternative is to
measure the permanent adduct, AFB1-FAPyr. Conversion of the major
AFB1-DNA adduct (AFB1-N7-Gua) to the more stable AFB1-FAPyr
(imidazole ring-opened guanine) is known to occur in the liver. Inter-
individual differences in levels of this permanent adduct suggest that
susceptibility to aflatoxin may also be genetically restricted (Lee et al.,
1989). With the development of monoclonal antibodies to specifically
quantify AFB1-FAPyr adducts, insight might be gained into the relation-
ship between infection with HBV, dietary exposure to AFB1 and liver
cancer. However the measurement of this adduct requires the taking of a
liver tissue sample, an invasive technique (Hsieh et al., 1988).
Ideally a laboratory assay is needed that will give assessment of long
term aflatoxin exposure in a sample obtained non-invasively. A highly
significant correlation between AFB1-N7-Gua excretion into urine and
AFB1 intake has already been shown, but the measurement of this para-
meter only reflects relatively short-term exposure. Groopman et al. (1987)
Overcoming aflatoxin contamination 535
have now shown a strong correlation between urinary excretion of
AFB1-N7-Gua adduct and AF-albumin ad ducts in the peripheral blood.
AFB1 binds in a dose-related manner to peripheral blood albumin in rats.
Upon repeated exposure, accumulation of binding occurs and the level of
albumin binding parallels the binding to liver DNA (Wild et al., 1990).
Because albumin has a half-life of about 20 days in humans, this adduct
may be a useful marker of aflatoxin exposure over a period of weeks or
months, with an accumulation of adducts to a level 30-fold higher than that
induced by a single exposure.
A direct ELISA of the AF-albumin adducts has been shown to give false
positive results. Hydrolysis of the sample gives an AF-lysine adduct.
Measurement of this by the same assay is much more sensitive, and can
detect 5 pg aflatoxin/mg albumin. This method can be used as an initial
screening test, followed by HPLC to confirm positive samples (Wild et al. ,
1990). Before blood AF-albumin levels can be extrapolated to liver DNA
damage, they must be assessed under experimental conditions which
reflect the human condition, especially when the liver is compromised by
environmental variants, such as HBV, alcohol and/or nutritional de-
ficiency. This is because studies with rats suggest that the two parameters
may be affected by alcohol intake. Until then, the measurement of the
albumin adduct will be useful only to identify populations at risk.
Measurement of the excreted DNA adducts will continue to be a useful
index for liver damage in exposed people (Wild et al., 1990).
13.10 OVERCOMING AFLATOXIN CONTAMINATION
In the past, the fairly high levels of aflatoxin often found in groundnut
meals restricted their use as a protein source in many animal diets. The
alternative was to divert this meal to fertilizer, which grossly underutilized
this valuable source of protein. Nowadays groundnuts are gaining even
more importance as a source of low-cost protein in the human diet,
especially in developing countries where aflatoxin contamination is most
severe. This stresses the need to eliminate or remove aflatoxins from
contaminated groundnuts.
Ideally the best control method is prevention - encouraging good farm
management in cultivation and storage practices. However, damage and
contamination still occur, which has led to various methods being
employed in the attempt to eliminate or lower the contamination to
acceptable levels for human and animal consumption. Aflatoxins have
proved to be highly resistant to various physical and chemical treatments.
A problem arises with the processing of the kernels into a variety of
products, e.g. oil, press and expeller cake (containing some residual oil)
and meal (solvent-extracted, therefore containing low oil levels). This
means that a particular method may be severely restricted by the nature of
the product to be detoxified (Feuell, 1966).
13.10.1 Removal of aflatoxin

Generally a relatively small number of kernels are contaminated and in
many cases physical separation is the ideal method. This is based on the
assumption that contaminated kernels are either discoloured or shrivelled.
Methods include manual sorting (Dickens and Weltz, 1968), mechanical
sorters or electronic scanning which examines each kernel separately,
scanning with a photoelectric cell which accepts or rejects them on the basis
of reflectance (Goldblatt, 1971).
Groundnut samples are currently graded as sound and mature if they
contain less than 2% damaged kernels, including discoloured, broken,
insect-infested or mould-damaged kernels. However, subsamples of a
given lot of these groundnuts may vary greatly in aflatoxin content, due to
the extreme variability in the degree of contamination in individual kernels
(Hodges et ai., 1964). Wrinkled nuts, thought to be caused by dampening
during storage, can account for 5% of a sample previously graded as
sound. While this is a relatively high level of contamination, the level of
toxin within these kernels is usually comparatively low. Conversely, a
second type of contamination is characterized by a low (0.24%) con-
centration of contaminated kernels but very high levels of toxin.
Disconcertingly, the majority of these kernels appear normal on the
exterior (Cucullu et ai., 1966).
When aflatoxin-contaminated nuts are pressed for oil, most of the
aflatoxin remains in the press-cake. The same occurs during oil extraction
with commercial hexane, with most remaining in the extracted by-product
meal. With the removal of the oil, the cake and meal have an enriched
protein content, which make them cheap protein sources for compounding
into animal feeds (Feuell, 1966). (The outbreak of Turkey X disease in the
UK in 1961 was caused by prepressed, hexane-extracted groundnut meal
containing a high level of aflatoxin.)
Several solvent extraction methods remove aflatoxins from contami-
nated oilseed meals. They have the advantage of being able to remove
essentially all aflatoxins without affecting the nutritive value or forming
adverse by-products. Polar solvents appear to be the most effective;
acetone and methanol have been shown to extract aflatoxins completely
from substrates. A binary solvent consisting of 90% acetone and 10%
water (w/w) , used for extracting aflatoxin from pre-extracted groundnut
meal of low oil content, gave up to 88% removal of aflatoxin. With this
system, an increased solvent temperature was shown to result in a higher
percentage reduction of aflatoxin. A tertiary solvent of 54% acetone, 44%
hexane and 2% water (w/w) was also developed for simultaneously extract-
ing oil and aflatoxin from prepressed groundnut cake (Gardner et ai.,
Overcoming aflatoxin contamination 537
1968). The water level in each system is critical. While it facilitates extrac-
tion, too much removes excessive amounts of water-soluble meal constitu-
ents and leads to processing difficulties. Dollear et ai. (1968) showed that a
binary solvent (90% acetone, 10% water) completely extracted aflatoxin,
leaving a meal with a chemical composition nearly identical to the original
starting material.
Aqueous isopropanol has also been shown to be an effective solvent for
the removal of aflatoxin from contaminated groundnut meals. Extraction
with six passes of 80% aqueous isopropanol at a minimum of 60°C resulted
in complete removal. Extraction with the isopropanol-water azeotrope
(87.7% isopropanol, w/w) was less effective, removing approximately 80%
of the aflatoxin (Rayner and Dollear, 1968).
Crude oils may contain various amounts of aflatoxin, depending on the
material used and the method of processing. Treatment with fuller's earth
and a hot alkali wash (conventional refining) has been shown to remove
essentially all aflatoxin present (Parker and Melnick, 1966). The aflatoxin
in crude oil appears to be in suspension, associated with cell debris carried
over mechanically, and not in true solution. This was confirmed by
Basappa and Sreenivasamurthy (1977) who showed that sedimentation by
centrifugation at 40 000 x g for 30 minutes removed a maximum of 65% of
aflatoxin. (This compared with a maximum 50% removed by filtration.)
The rest (35%) remained in solution in the supernatant. Studies with
different adsorbants showed that activated fuller's earth (2%) increased
sedimentation of aflatoxin to 92.5% at 10 000 x g for 30 minutes.
Alternatively the use of casein or arecanut powder (35% total tannins), at
10% w/w, reduced the aflatoxin by 92% and 86% respectively but gave
objectionable colour and flavour to the oil (Dwarakanath et ai., 1969).
Another recent study has shown hydrated sodium calcium aluminosili-
cate (HSCAS), used as an anti-caking agent in animal feed, to be a high
affinity adsorbant for aflatoxin (Harvey et ai., 1989; Kubena et al., 1990;
Pettit et ai., 1990). Studies showed that more than 80% aflatoxin was
bound to HSCAS. Less than 10% of this was subsequently extracted,
indicative of the strong bonds that are formed. Kubena et ai. (1990) found
that the growth inhibitory effects of AFB1 were significantly diminished
when 7.5 mg/kg AFB1 was added to poultry feed containing 0.5% HSCAS.
Pigs fed a diet containing 3mg/kg and 0.5% HSCAS showed no liver
lesions (Harvey et ai., 1989). These findings suggest that HSCAS can
reduce the bioavailability and therefore toxicity in growing animals. As
such, it presents a novel approach to preventative management of the
problem of aflatoxin contamination in such feeds.
13.10.2 Destruction of aflatoxin

Destruction of aflatoxin, whether by heat, chemical or biological methods,
is another option. With such methods, however, there is always the
possibility of diminished nutritive value or unwanted residues. Proteins are
susceptible to denaturation; groundnut protein is also lysine-deficient,
even before the terminal amine group of this amino acid is attacked by
reactive chemicals.
Aflatoxin is very stable to heat at temperatures below 100°C. Several
studies have been carried out on the effects of dry- and oil-roasting on
groundnut kernels. Lee et al. (1969) showed the average reduction of
aflatoxin ranged from 43 to 83%, depending on time and temperature
conditions and the level of toxin in the raw kernels. They reported an
overall reduction of 65% in AFBI for oil roasting compared with 69% for
dry roasting, with the percentage reduction being highest at the higher
aflatoxin contamination levels.
In a similar experiment, Pluyer et al. (1987) showed that oven-roasting at
150°C for 30 minutes caused a 30-45% reduction of AFB1. They also
showed that microwaving at 0.7 kW for 8.5 minutes gave a similar
reduction in aflatoxin levels. (Another group reported a 95% reduction
after 4 minutes at 6 kW, but to achieve this reduction at lower. energy levels
would lead to over-roasting the kernels.) Unlike Lee's group (1969), no
correlation was found between the percentage destruction and the levels of
aflatoxin in the samples.
Autoc\aving wet meals reduces aflatoxin content but this can take up to 4
hours (Coomes et al., 1966). Codifer et al. (1976) reduced this to 1 hour by
the addition of 1.5% formaldehyde and 2% calcium hydroxide, but the
nutritional value of the final product was in doubt.
Many chemicals have been screened for their potential to destroy
aflatoxin (Feuell, 1966; Sreenivasamurthy et al., 1967; Dollear et al.,
1968; Natarajan et al., 1975b). Under alkaline conditions the lactone
ring common to AFB1 and AFB2 hydrolyses to the hydroxy acid
form, which is then susceptible to oxidation. Ammonia gas or
ammonium hydroxide detoxify by alkaline hydrolysis of the lactone
moiety. Vesonder et al. (1975) showed apparent detoxification with
ammonium hydroxide at room temperature for 21 hours. Subsequent
treatment with acid restored the AFBI to its toxic form, which
demonstrates the need for caution when assessing potential chemicals
to process meals.
It is now known that several compounds are formed during the ammo-
niation process. If the process is carried out under mild conditions, then
the reversibility demonstrated by Vesonder et al. (1975) can occur.
However, when the reaction is allowed to proceed further, the compounds
formed do not revert back to aflatoxin (Park et al., 1988). Since ammonia-
tion has been shown to reduce aflatoxin levels to less than 1%, it is
becoming the method of choice. Ammonium hydroxide or gaseous
ammonia can be used at either high temperature and pressure or low
temperature and pressure (Dollear et al., 1968; Gardner et al., 1971;
Viroben et al., 1978; Delort-Laval et al., 1980; Park et al., 1988). Reaction
Conclusions 539
products formed depend on the treatment and conditions used (Lee and
Cucullu, 1978; Park etai., 1988).
The ammoniation process reduces the amount of soluble nitrogen in the
meal. Cystine undergoes partial destruction, which can be counterbalanced
by the addition of synthetic methionine in animal diets. Protein efficiency
ratios (PER) are generally lower, probably due to the decreased nitrogen
solubility and available lysine levels (Gardner et ai., 1971; Viroben et ai.,
1978; Delort-Laval et ai., 1980). However, Park et ai. (1988) showed
production parameters, e.g. milk and egg quality, to be significantly better
or not adversely affected by the treatment. The total nitrogen content of
the meals increased, resulting in positive weight gain and feed efficiency
values in ruminants. Park et al. (1988) reported that either no effect or
decreased weight gain or feed efficiency values were obtained with simple-
stomached animals, which correlates with Dollear et ai. (1968) who
reported that treated meals gave lower weight gains with rats.
Frayssinet and Lafarge-Frayssinet (1990) showed the efficacy of ammo-
niation in a long-term feeding study with rats. The aflatoxin content
decreased proportionately with the increased pressure of application. The
degradation products of the reaction showed no carcinogenicity; the pro-
cess did not produce any new toxin and no reversibility was observed.
There are conflicting reports as to the effectiveness of UV radiation in
destroying the toxin (Feuell, 1966; Goldblatt, 1971). Shantha (1989)
showed that the exposure of contaminated groundnut oil to bright sunlight
for 15-30 minutes completely destroyed the toxin. Exposing it to UV light
was far less effective. Gamma radiation at acceptable safety levels has been
shown to be ineffective (Feuell, 1966; Chiou et ai., 1990). Van Dyck et ai.
(1982) had success with ionizing radiation in aqueous solutions, but the
dose required to detoxify the aflatoxin (> 10 kGy) was outside the permiss-
ible range for the irradiation of foodstuffs. Patel et ai. (1989) expanded this
work by demonstrating the synergistic effect of hydrogen peroxide and
gamma radiation to degrade higher concentrations of aflatoxins at lower
doses of radiation. This followed on from work by Sreenivasamurthy et ai.
(1967), who demonstrated the effectiveness of 6% hydrogen peroxide
under alkaline conditions in destroying 85% of the toxin. Together,
gamma radiation and hydrogen peroxide produce a higher amount of free
radicals in aqueous solution than individually. This method is suited to
groundnut kernels and has the advantage over ammoniation of being less
time-consuming and a less expensive process. However, animal toxicity
studies are required before commercial application is possible.
13.11 CONCLUSIONS
Aflatoxin contamination of groundnuts is the most significant econ-

omic problem facing the groundnut industry. Once groundnuts are
contaminated, there are severe limitations placed on the removal of
aflatoxin. For example, while solvent extraction may be applicable for the
treatment of groundnut meals, it is not the method of choice for whole
kernels. Apart from these individual limitations, there is a general one of
cost; a detoxification step involving additional handling and processing
would inevitably raise the final cost of the commodity. There is also the
possible nutritive loss (Feuell, 1966; Goldblatt, 1971). Therefore it has to
be stressed that prevention of contamination is still the best approach.
The marketable commodities produced from groundnuts, including
edible oil, expeller meal, oil cake, confectionery products and peanut
butter, each represent a different level in the trade chain. There is the
potential for reduced export earnings and loss at each level if aflatoxin
contamination is detected above certain limits.
An enormous range of aflatoxin concentrations in groundnut products
has been reported between countries. It must be borne in mind that this
could be due to variation in the sensitivity of the analytical methods
employed rather than the true levels of contamination.
Many countries have now set limits for AFB1 and total aflatoxin in
foodstuffs and feed, as well as allowable limits of AFM1 in dairy products
(Van Egmond, 1989). Data collected over the last quarter of a century
indicate that the risk from consumption of aflatoxin-contaminated ground-
nuts is likely to be more to animals than to humans (Bhat, 1989). Limits on
animal feed not only protect the health and productivity of the animals
consuming groundnut-containing feeds but also prevent secondary trans-
mission to humans via the food chain.
The major problem lies with developing countries. The environmental
conditions in these countries are often ideal for aflatoxin production. This,
combined with poor management practices during production, harvest,
handling and storage, must affect the degree of contamination. Farmers
will surely sell their crops when heavier (and therefore insufficiently dry) if
it means better financial returns. In many places up to 80% of the agricul-
tural produce is consumed at home or at the village level, without entering
the organized market chain (Bhat, 1989). This means that there is no
dilution effect, and these people and livestock can be exposed to large and
prolonged doses of aflatoxin. It is these same people (and their animals)
who are often malnourished and therefore more susceptible to the toxin
and its immunosuppressive effect.
ACKNOWLEDGEMENTS
The assistance of the staff of the Lincoln University Library in obtaining

many of the references used in this work is gratefully acknowledged.
References 541
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CHAPTER 14
Groundnut breeding
T.G. Isleib~ Ie. Wynne and S.N. Nigam
14.1 INTRODUCTION
Groundnuts (Arachis hypogaea L.) are grown throughout the tropical and
warm temperate regions of the world, with commercial production princi-
pally between latitudes 40° Nand 40° S. Leading producing nations are
India (33.4% of global production), China (27.8%), USA (9.3%), Senegal
(4.2%), Indonesia (4.2%), Nigeria (3.3%), Myanmar (3.0%), Sudan
(2.7%) and Argentina (2.0%). Clearly, the crop is grown in several
agroecological systems and under numerous socioeconomic environments.
Yield of groundnuts is often low due to diseases and insects, unpredictable
and variable rainfall, inability to apply improved agronomic practices and
production technology, lack of cultivars adapted to local conditions, low
financial inputs, lack of small-scale farm implements, and lack of the
infrastructure required to supply quality seed of improved cultivars (Nigam
et al., 1991).
The primary objectives of groundnut breeders are to develop cultivars
with high yield potential, adaptation to specific environments and pro-
duction systems, resistance or tolerance to environmental stresses and
resistance to diseases and insects. Because groundnuts are grown under
many different cropping systems across a wide array of agroecological
conditions, the specific objectives of breeding programmes vary consider-
ably. Breeding is a continuing process as the crop is introduced to new
environments and production systems, as market demands change, and as
disease and insect pest populations shift in reaction to deployment of new
cultivars. As the primary constraints to production are overcome by new
cultivars and production practices, breeding for improved flavour and
quality desired by processors and consumers becomes more important
(B unting et al., 1985).
Variability in groundnut germplasm 553
TABLE 14.1 Botanical division of A. hypogaea L.
A. hypogaea L. Characteristics Secondary centres of

diversity
subsp. hypogaea No flowering on main stem

Alternate branching
vaL hypogaea Two seeds per pod Bolivian
Amazonian
var. hirsuta Kohler Long mainstem Peruvian
Three or more seeds psr pod
subsp. fastigiata Flowering on mainstem
Waldron Sequential branching
vaL fastigiata Limited vegetative branching Goias and Minas Gerais
Three or more seeds per pod Guaranian
Goias and Minas Gerais
Peruvian
North-east Brazil
vaL vulgaris Harz Short runs of reproductive Goias and Minas Gerais
branches
Two seeds per pod Guaranian
Peruvian
14.2 VARIABILITY IN GROUNDNUT GERMPLASM
In order to develop cultivars with traits that overcome the constraints

peculiar to a specific environment, there must be sufficient genetic
variation to allow selection for desired traits. Assertions by American
researchers regarding the paucity of genetic variation in groundnut
referred to specific economically important characters of the extant culti-
vars and breeding stocks within market classes in the USA (Gregory et ai.,
1973). More recently, molecular analyses have not detected significant
amounts of variability in allozymes (Grieshammer and Wynne, 1990),
restriction fragment length polymorphisms (Kochert et ai., 1991) or DNA
fragments amplified by polymerase chain reaction (Halward et ai., 1992) in
cultivated germplasm of broadly diverse origin.
The most commonly used botanical division of A. hypogaea into subspe-
cies and varieties is that of Krapovickas (1968), based on patterns of
reproductive and vegetative branching and on pod morphology as summar-
ized in Table 14.l.
Because of strong local preferences for particular pod and seed charac-
teristics, early breeders of ground nut often worked with limited numbers
of parents possessing attributes acceptable to local consumers or pro-
cessors. In the USA, market classes of groundnut roughly follow the
554 Groundnut breeding
botanical divisions of the cultivated species with the following exceptions.
Firstly, the runner and virginia market classes are commonly equated with
var. hypogaea, but do not have purely hypogaea ancestry. Both have had
substantial introgression of fastigiate germplasm, primarily from spanish
ancestors (var. vulgaris), in the course of plant improvement through
breeding (Isleib and Wynne, 1992). Spanish parents were used to increase
oil content, shorten maturity and increase the harvest index of the crop.
The most common runner and virginia groundnut cultivars in the USA
have 0-50% fastigiate ancestry (Table 14.2) and average 35%. Secondly,
groundnuts of var. hirsuta are not represented in any market class, nor has
hirsuta germplasm been used in the development of any released cultivars
or registered germplasm to date. Hirsuta types are extremely rare in the
USA national collection and in the global collection maintained at the
International Crops Research Institute for the Semi-Arid Tropics (ICRI-
SA T) in Andhra Pradesh, India. Chinese scientists report that 'peruvian-
type' groundnuts are grown commonly in their central production region,
but it is not clear whether the peruvian type mentioned is hirsuta or
Jastigiata with the typical peruvian pod configuration. Hirsuta groundnuts
remain a garden crop in north-western South America and in Mexico
(D. Williams, personal communication), and collection efforts should be
focused there.
It was not until the late 1970s that the extent of natural genetic variability
available to groundnut breeders was fully appreciated and widely recog-
nized (Norden, 1980). Collections of cultivated groundnuts were con-
sidered extensive by the mid 1970s (Banks, 1976). Many of these and other
accessions in germplasm collections in the USA or at ICRISAT since 1976
were obtained from expeditions made to South America, the centre of
origin and diversity for groundnuts, under the sponsorship of the US
Department of Agriculture (USDA) and the International Board for Plant
Genetic Resources (IBPGR) in co-operation with state experiment
stations in the USA and several other countries. The most important
expeditions were those of Archer in 1936; Stephens and Hartley in 1947-
48; Gregory, Krapovickas, Pietrarelli and others in 1959, 1961, and 1967;
Hammons, Langford, Krapovickas, Pietrarelli and others in 1968; and
Gregory, Banks, Simpson, Krapovickas, Pietrarelli and others in 1976,
1977, 1979 and 1980 (Wynne and Gregory, 1981) as well as those of
IBPGR/CENARGEN/ICRISAT teams made during 1989 (Simpson,
1990). Collection of South American genetic resources continues today
with particular emphasis on the wild species native to areas of Brazil
undergoing rapid development. Much of the germplasm from Africa, an
important centre of secondary variation (Gibbons et al., 1972), was intro-
duced into the United States by Smartt in 1959 (Wynne and Gregory,
1981). These accessions are maintained by the Southern Regional Plant
Introduction Station at Experiment, Georgia. The most extensive collec-
tion of cultivated groundnut germplasm is now maintained by ICRISAT.
The Genetic Resources Unit there maintains a global collection of more
than 12000 accessions (Nigam et al., 1991). In addition to the cultivated
germplasm, there are more than 70 wild species of Arachis (Stalker and
Moss, 1987). Some of the wild species have direct value as forages (Prine et
al., 1981) but for the most part they constitute a genetic reservoir of useful
characteristics for the improvement of the cultivated groundnut, notably
with respect to host-plant resistance to diseases and insects but perhaps
also for agronomic traits (Guok et al., 1986).
Although it was widely recognized that there was tremendous morpho-
logical variation among the accessions of cultivated groundnuts, it has only
been during the last decade that the extent of the desirable variation has
been demonstrated (Wynne et al., 1991). As late as 1973, the widely held
view was that there were many defects, with respect to the requirements of
man, in the genetic composition of groundnuts (Gregory et al., 1973). This
view resulted from inadequate collection and evaluation of the germplasm
of groundnuts (Wynne and Halward, 1989b). Systematic and extensive
screening of the cultivated germplasm was not practised until ICRISAT
adopted groundnut as a mandate crop in 1976 (Nigam et al., 1991).
Several groundnut breeders had large collections of groundnuts by the
1970s but these were inadequately evaluated due to limited funds and
personnel (Norden, 1980). Although many of the efforts to evaluate
groundnut germplasm have not been systematic or exhaustive, a large
number of accessions of cultivated groundnuts have been identified that
contain desirable variation for yield, fruit size, morphological traits, toler-
ance to environmental stresses, disease and insect resistances, and seed
composition.
In many developing nations, groundnut's primary use is as an oilseed
either for domestic use or for export. Other countries view groundnut
primarily as a food crop. In either case, the composition of groundnut has
become an issue of increasing importance in the past ten years. For use as
an oilseed, the market demands groundnuts with high oil content and good
storability. Where groundnut is used as a food, whether as whole or
processed seeds, attributes important to the consumer such as flavour,
protein quality and shelf life have long been primary concerns of the
groundnut marketing and processing industries. Shelf life is largely a
function of the time required for auto-oxidation of linoleic fatty acid in the
oil fraction of the seed to produce a characteristic rancid flavour. Increased
use of oxygen-permeable packaging materials by groundnut processors has
increased the need to extend shelf life through genetic improvement of the
seed itself. Oleic acid, the 18-carbon mono-unsaturated (18:1) precursor to
linoleic acid (18:2), is less reactive with oxygen and is therefore more
desirable in the fatty acid profile of groundnut oil. A commonly used index
of the storability of groundnut is the ratio of oleic to linoleic fatty acids
(OIL ratio). This ratio ranges in value from under 1 to over 2 in cultivars
used in the USA (Brown et al., 1975; Ahmed and Young, 1982). OIL ratio
TABLE 14.2 Fastigiate ancestry of runner- and virginia-type groundnut cultivars currently or formerly grown in the United States
Cultivar Market class Share of US Specific fastigiate ancestry Total fastigiate

certified area l ancestry
Dixie Runner Runner 0% 50% Small White Spanish 3x-1 50%

Early Runner Runner 0% 50% Small White Spanish 3x-2 50%
Florunner Runner 47.4% (61.8% ) 37.5% Small White Spanish 3x-2 50%
12.5% Spanish 18-38
GK7 Runner 7.0% (9.1%) 15.625% Small White Spanish 3x-2 31.25%
15.625% Spanish 18-38
MARC I Runner 0% 1.5625% McSpan Spanish 40.625%
(New release) 3.125% Pearl
6.25% Small White Spanish 3x-1
7.8125% Spanish 18-38
Okrun Runner 3.7% (4.8%) 50% Argentine 75%
6.25% Spanish 18-38
Southern Runner Runner 2.2% (2.9%) 18.75% Small White Spanish 3x-2 25%
6.25% Spanish 18-38
Sunrunner Runner 3.5% (4.5%) 25% Small White Spanish 3x-2 37.5%
12.5% Spanish 18-38
TABLE 14.2 Cant.
Cultivar Market class Share of US Specific fastigiate ancestry Total fastigiate

certified area I ancestry
Florigiant Virginia 0.3% (1.9% ) 25% Small White Spanish 3x-2 37.5%
12.5% Spanish 18-38
NC2 Virginia 0% 25% Spanish 18-38 25%
NC5 Virginia 0% 25% Improved Spanish 2B 25%
NC6 Virginia 1.3% (8.0% ) 0%
NC7 Virginia 5.2% (33.4%) 6.25% Improved Spanish 2B 43.75%
25% Small White Spanish 3x-2
12.5% Spanish 18-38
NC9 Virginia 3.6% (23.0%) 12.5% Small White Spanish 3x-2 31.25%
18.75% Spanish 18-38
NC lOC Virginia 2.2% (13.9%) 6.25% Improved Spanish 2B 37.5%
12.5% Spanish 18-38
NC-Vll Virginia 3.0% (19.1%) 6.25% Improved Spanish 2B 50%
6.25% Spanish 18-38
25% PI 337396 (var. JastigiataI)
VA-C 92R Virginia 0% 6.25% Improved Spanish 2B 28.125%
(New release) 12.5% Small White Spanish 3x-2
9.375% Spanish 18-38
[Proportion of total area certified and (in parentheses) proportion of market class area certified.
is commonly used as a criterion for release of new cultivars in the USA
with high values viewed as desirable by the groundnut industry. It is
interesting to note that this demand for mono- rather than di-unsaturated
fat in groundnut contradicts the general demand for less saturated oils in
components of human diets. Norden et al. (1987) reported a groundnut
variant with an extremely high OIL ratio greater than 30. Moore and
Knauft (1989) determined that this trait was governed by two recessive
genes and is therefore easily transferable to existing cultivars through
backcrossing.
Recently, concern over the high contribution of fats to daily caloric
intake by the populations of industrialized nations has created demand for
low- or reduced-fat foods. The range of oil content in cultivars in the USA
is 43.6-55.5% (Norden et al., 1982). Among over 6000 groundnut acces-
sions evaluated at ICRISAT, the range was 31.8-55.0% (ICRISAT, per-
sonal communication). This range is too high to permit reference to
groundnut, even at the lower extreme of the distribution of fat contents, as
a low-fat food, but it is clear that selection for reduced oil content should
be effective.
14.2.1 Foliar fungal pathogens

Much of the screening of the groundnut germplasm for desirable variation
during recent years has emphasized biotic stresses. This work was recently
reviewed (Nigam et al., 1991; Wynne et al., 1991). Three foliar fungal
diseases -late leaf spot [Phaeoisariopsis personata (Mycosphaerella berke
leyi)], early leaf spot [Cercospora arachidicola (Mycosphaerella arachidis)]
and rust (Puccinia arachidis) - are the most widely distributed and econ-
omically important diseases of groundnut. They are common wherever
groundnuts are growl? but they vary in incidence and severity among
locations and years. Each disease alone can cause severe damage but yield
losses are generally increased when they occur together. For example,
rust and late leaf spot together can cause up to 70% yield loss in India
(Subrahmanyam et al., 1984). These diseases also affect seed grade
adversely and they markedly reduce haulm yields - an effect that is of
particular importance in those regions of the semi-arid tropics where small
farmers maintain significant numbers of livestock.
Effective field screening methods have been developed for use in areas
where natural disease pressure is high or where such pressure can be
artificially induced. At the ICRISAT Center, field screening with infector
rows is used to challenge host plants in a worst-case situation (Subrahma-
nyam et al., 1982a). Genotypes and breeding populations to be screened
are planted in a nursery together with rows of highly susceptible cultivars
arranged systematically throughout the nursery. The ratio of test and
infector rows varies from season to season and location to location. A
mixture of short- and long-season susceptible cultivars is used to ensure
inoculum supply for a longer period. Plants in infector rows are inoculated
with spore suspensions to enhance disease development. This procedure is
most successful if infector rows are inoculated in the evening immediately
following overhead irrigation. Potted 'spreader' plants heavily infected
with rust are also placed systematically throughout the field to provide
another source of inoculum. The nursery may be irrigated by overhead
sprinklers until harvest as required by climatic conditions.
Disease reaction on test plants is scored using a nine-point scale (Subrah-
manyam et al., 1982a, b). Disease scores are recorded about 10 days before
harvest in preliminary screening and at several growth stages in advanced
screening and other studies. These techniques are useful for grouping lines
into resistant and susceptible classes but not for identifying moderate levels
of resistance. Germplasm and advanced breeding lines can also be
screened in the glasshouse using potted plants or in the laboratory using
detached leaves to measure components of disease resistance such as latent
period, lesion number, lesion size and sporulation rate.
Screening methods similar to those used for rust are also used for late
leaf spot using plots of test genotypes interspersed at regular intervals with
susceptible infector rows inoculated with late leaf spot spores. Additional
inoculum is provided by scattering on the infector rows leaf debris col-
lected from infected plants in the previous season.
In the USA, field methods for identifying moderate resistance to leaf
spots generally require isolation of test genotypes from one another to
minimize the effect of adjacent plots. In North Carolina, isolation has been
accomplished with border rows of non-host crop species such as maize (Zea
mays L.), soybean (Glycine max [L.] Merr.) or cotton (Gossypium hirsu-
tum L.) with eight border rows at 90 cm spacing between adjacent plots and
6-7 m of border between plots occupying the same rows in the field. Each
plot is inoculated with spore suspension to ensure the presence of the
pathogen, and disease progress is monitored after inoculation. Defoliation
of the mainstem is the primary criterion of resistance and is expressed as a
proportion or percentage of nodes defoliated. Isolation of the plots reduces
the influence of neighbouring susceptible plants on accessions with partial
resistance.
Sources of resistance to rust were reported by Bromfield and Cevario
(1970). Hammons (1977) summarized the screening of groundnuts for rust
resistance and concluded that resistant sources originated from three
sources: Tarapoto (PIs 259747, 341879, 350680, 381622, and 405132),
Israeli line 136 (PIs 298115 and 315608), and DHT 200 (PI 314817).
Tarapoto and DHT 200 both originated in Peru. ICRISAT has screened
more than 12000 accessions of groundnut for rust resistance in the field,
using infector rows to develop disease pressure (Subrahmanyam and
McDonald, 1983),and 124 lines have been found with rust resistance
(Nigam et al., 1991). These include 14 rust-resistant lines released jointly
by USDA and ICRISAT (Subrahmanyam and McDonald, 1983). In addi-
tion, several wild Arachis species and their interspecific derivatives with
cultivated groundnuts have been screened for resistance to rust under both
field and laboratory environments (Subrahmanyam et ai., 1983c). The rust
resistance identified in the cultigen is of a 'slow-rusting' type. Resistant
lines exhibit increased incubation period, decreased infection frequency
and reduced pustule size, spore production and spore germinability
(Subrahmanyam et ai., 1983a, b). Many wild Arachis species, and lines
derived from their hybridization with the cultigen, have been screened for
resistance to rust under field and laboratory conditions. Accessions of
several species were found to be immune to rust: A. batizocoi (PI 298639,
PI 338312), A. duranensis (PI 219823), A. cardenasii (PI 262141), A.
chacoensis (PI 276235), A. pusilla (PI 338449), A. villosa (PI 210554) and
A. correntina (PI 331194) among others (Subrahmanyam et ai., 1983c).
Most of the interspecific derivatives showed a high degree of resistance to
rust. They had small and slightly depressed uredinia that did not rupture to
release the comparatively few uredospores produced.
Screening for resistance to the leaf spots caused by C. arachidicoia and
P. personata has been extensive in recent years. Several sources of resist-
ance to both early and late leaf spots have been reported (Foster et ai.,
1980, 1981; Gorbet et ai., 1982; Hassan and Beute, 1977; Melouk et ai.,
1984; Subrahmanyam et ai., 1985). Screening for late leaf spot resistance
has been most extensive at ICRISAT where the 12000 or more genotypes
screened for rust have also been screened for late leaf spot. Fifty-three
accessions of A. hypogaea have now been identified with documented
resistance to late leaf spot (Nigam et ai., 1991) and 29 of these 53 lines are
also resistant to rust (Table 14.3). Resistance to late leaf spot operates
through much the same mechanisms as resistance to rust (Subrahmanyam
et ai., 1982b).
Among the many accessions of wild Arachis species tested at ICRISAT
Center, A. chacoense (PI 276325), A. cardenasii (PI 262141) and A.
stenosperma (PI 338280) of section Arachis combined cross-compatibility
with the cultigen and immunity or high resistance to the pathogen. Highly
resistant wild species from other sections included A. repens, A. appressi-
piia, A. paraguariensis, A. villosulicarpa, A. hagenbeckii and A. giabrata
(Subrahmanyam et ai., 1985).
Several lines, including NC 3033, NC 5, PI 270806, GP-NC 343, PI
109839, PI 259747 and PI 350680, have been shown to possess epidemiolo-
gical components of rate-reducing resistance to early leaf spot in the USA
(Foster et ai., 1981; Green and Wynne, 1987; Hassan and Beute, 1977;
Sowell et ai., 1976). Some of these lines (NC 3033, PI 270806, PI 259747
and PI 350680) did not show resistance in India or Malawi when infector-
row inoculation techniques were used (Nigam, 1987; ICRISAT, 1984).
Because early leaf spot does not usually occur readily in the field at
Patancheru, screening has been less extensive at ICRISAT than for either
rust or late leaf spot, but incidences of heavy disease at ICRISAT Center in
1983 and 1987 were utilized to screen germplasm already planted in the
field. Also in 1987, screening for early leaf spot resistance on a limited scale
was initiated by ICRISAT in Pantnagar, India, where C. arachidicola
occurs more regularly. Of 3000 genotypes screened for early leaf spot,
several showed moderate levels of field resistance at both locations
(Waliyar et al., 1989):
ICG 2711 (NC 5)
ICG 6709 (NC Ac 16163)
ICG 7291 (PI 262128)
ICG 7406 (PI 262121)
ICG 7630
ICG 7892 (PI 393527-B)
ICG 9990.
In Malawi, screening for early leaf spot resistance has not identified
significant sources of resistance. More than 1000 selected germplasm lines
of the cultigen have been screened individually but none showed any
appreciable level of resistance to the disease. In 1986-87, 'bulk' testing was
utilized to evaluate a large number of lines: 110 bulk populations were
constructed by compositing five seeds from each of 100 lines. All lines in a
given bulk population shared a common botanical variety. This method
allowed representation of 11 000 lines in the screening although the identi-
ties of individual lines were lost. Only two bulks had a few plants which
merited further testing. In the 1987-88 season, component lines of the two
bulks were planted separately and scored for the disease. Only three
germplasm lines - ICG 50, ICG 84 and ICG 11282 - were retained for
further testing. Other lines that retained a higher than usual proportion of
foliage despite heavy disease pressure (ICRISAT, 1986) were:
ICGM 189 (ICG 5216, PI 262087)
ICGM 197 (ICG 6012, NC Ac 16142, PI 262093)
ICGM 281 (ICG 8515)
ICGM 284 (ICG 8521)
ICGM 285 (ICG 8522)
ICGM 286 (ICG 8523)
ICGM 292 (ICG 8529)
ICGM 300 (ICG 8569, NC Ac 868, PI 119072)
ICGM 473 (ICG 3431)
ICGM 500 (ICG 3150)
ICGM 525 (ICG 6151).
Thirty-five lines reported to have resistance to early leaf spot at ICRISAT
Center were not resistant in Malawi (ICRISAT, 1989).
Several wild species of Arachis, including A. cardenasii, have been
reported to be resistant to early leaf spot; however, only Arachis species
30003 has shown consistent resistance when tested in Malawi using
TABLE 14.3 Sources of resistance to both rust (Puccinia arachidis) and late leaf spot (Phaeoisariopsis personatum) available at fCRfSAT
Centre in 1989
Disease score 2
ICG No. Identity Botanical variety Seed colour Origin Rust Late leaf spot
1703 NC Ac 17127 fastigiata Variegated Peru 4.7 5.0

1707 NC Ac 17132 fastigiata Purple Peru 4.0 4.0
1710 NC Ac 17135 fastigiata Dark purple Peru 4.0 4.0
2716 EC 76446 (292) fastigiata Dark purple Uganda 3.3 4.7
3527 USA 63 fastigiata Purple USA 4.7 4.7
4747 PI 259747 fastigiata Purple Peru 3.7 4.0

4995 NC Ac 17506 fastigiata Purple Peru 4.3 4.3
6022 NC Ac 927 fastigiata Purple Sudan 4.0 4.0
6330 PI 270806 hypogaea Tan Zimbabwe 2.1 3.3
6340 PI 350680 fastigiata Dark purple Honduras 3.0 4.0
7013 NC Ac 17133(RF) fastigiata Dark purple India 3.3 4.0

7881 PI 215696 fastigiata Dark purple Peru 4.3 3.7
7884 PI 341879 fastigiata Purple Israel 3.0 3.7
7885 PI 381622 fastigiata Purple Honduras 3.0 4.3
7886 PI 390593 fastigiata Tan Peru 4.7 3.3
7894 PI 393641 fastigiata Variegated Peru 4.0 4.7

TABLE 14.3 Cant.
Disease score 2
leG No. 1 Identity Botanical variety Seed colour Origin Rust Late leaf spot


10889 PI 476016 fastigiata Red Peru 3.3 4.3
10936 PI 476168 fastigiata Dark purple Peru 4.3 4.0
10941 PI 476174 fastigiata Light purple Peru 4.7 4.7

11485 fastigiata Light purple Peru 5.0 3.7
Susceptible check cultivars:

221 TMV2 vulgaris Tan India 8.3 8.0
799 Robut 33-1 hypogaea Tan India 7.7 7.3
1 ICRISA T ground nut accession number.

2 Scored on a modified 9-point disease scale where 1 = 0%, 2 = 1 to 5%, 3 = 6 to 10%, 4 = 11 to 20%, 5 :.= 21 to 30%, 6 = 31 to 40%, 7 = 41 to 60%,
8 = 61 to 80% and 9 = 81 to 100% damage to foliage (ICRISAT Center, rainy season 1989).
TABLE 14.4 Reaction of some groundnut germplasm lines with resistance to early
(Cercospora arachidicola) and late leaf spot (phaeoisariopsis personatum) and rust
(puccini a arachidis), ICRISA T Center, rainy season 1987
Disease reaction 1
Entry Original name Early Late Rust

leaf spot leaf spot
ICG 1703 NC Ac 17127 4.7 5.0 4.7

ICG 6284 NCAc17500 5.0 7.0 3.3
ICG 7340 198/66 Coli 182 5.7 5.1 2.7
ICG 9294 58-295 5.1 6.0 2.7
ICG 10010 PI 476143 5.7 5.1 4.1
ICG 10040 PI 476176 (SPZ 451) 5.0 4.7 3.7
ICG 10900 PI 476033 5.3 4.7 4.1
ICG 10946 PI 476176 5.0 6.0 4.1
Susceptible controls
ICG 799 Kadiri 3 (Robut 33-1) 8.0 7.0 7.0
ICG 221 TMV2 8.0 8.0 8.0
Mean (n=500) 6.9 6.5 4.9

Standard error2 ±0.48 ±0.7 ±1.1
CV(%) 7.0 11 22
1 Mean of 3 plots, each 2 4-m rows, rated on a 1-9 scale where l=no disease and 9=50-100%
foliar destruction.
2 Standard error and CV calculated on the basis of all 500 genotypes tested.
infector-row inoculation techniques (ICRISAT, 1989). Among other

species, A. chacoensis and A. sp. 30085 showed high promise in the first
year of screening but were susceptible in subsequent tests. A. stenosperma
was found to be highly susceptible in Malawi (ICRISAT, 1988), contrary to
reports from the USA. Several interspecific derivatives were found to
retain more foliage than the susceptible control cultivar.
Table 14.4 shows that eight lines of groundnuts with moderate to high
levels of resistance to all three foliar diseases - rust, early leaf spot and late
leaf spot - have been identified (ICRISAT, 1988; Waliyar etal., 1989). The
rust and late leaf spot reactions of most accessions are stable over a wide
range of geographic locations. Only for NC Ac 17090 and PI 298115 has
variation in rust reaction been observed across locations. Reaction to early
leaf spot has exhibited greater variation across locations. The eight lines in
Table 14.4 are potentially the most useful parental lines available, since the
foliar diseases generally occur in combination.
The genetics of resistance to these three diseases is not well understood.
Bromfield and Bailey (1972) first reported that resistance to rust in the
cultigen was controlled by two recessive genes. However, Nigam et al.
(1980) found continuous variation in the progeny of crosses among rust-
resistant FESR lines (Bailey et at., 1973) and suspected that rust resistance,
though recessive in nature, might be governed by more than two genes. In
generation means analysis of resistant-by-susceptible crosses, Reddy et al.
(1987) found additive, additive-by-additive and additive-by-dominance
effects for rust resistance. In some diploid wild Arachis species, resistance
appeared to be partially dominant (Singh et at., 1984).
Nevill (1982) studied five F2 progenies from crosses between two resist-
ant and three susceptible cuItivars for components of resistance to late leaf
spot in detached leaf tests. To account for the observed distribution of
phenotypic values in the F 2, he postulated a five-locus polygenic system
assuming resistance to be completely recessive. Non-additive gene action
was concluded to be extremely important but its nature could not be
elucidated due to the omission of the Fl generation from the study.
14.2.2 Viral pathogens

Variation for resistance to several virus diseases has been reported in
groundnut (Nigam et al., 1991). The crop is host to several viruses but only
a few are considered economically important. These include groundnut
rosette (GRV) in Africa, bud necrosis (BNV) in India, tomato spotted wilt
(TSWV) in the USA, peanut mottle (PMV) worldwide, peanut stripe
(PStV) in east and south-east Asia, and peanut clump (PCV) in West
Africa and India. Laboratory and field screening techniques have been
developed for all these virus diseases. Resistance to rosette virus was
discovered in local land races in Burkina Faso in the 1950s (de Berchoux,
1958, 1960). Of seven wild species of Arachis screened in an SADCC-
ICRISAT regional groundnut project, two species (A. sp. 30003 and A. sp.
30017) remained symptom-free throughout the season. The apparent
immunity of A. sp. 30003 to rosette and its high resistance to early leaf spot
suggest that efforts to use this species should be emphasized (Bock, 1989).
Several groundnut accessions with consistently low symptoms of bud
necrosis have been identified at ICRISAT, including:
C102
C121
C136
GP-NC 343
NC Ac 2232
NC Ac2242
NC Ac 17888
ICGV 86029
ICGV 86031.
Only ICGV 86029 and 86031 showed tolerance to the virus (Nigam et at.,
1991). Southern Runner, a cultivar with resistance to late leaf spot, has
shown fewer symptoms of tomato spotted wilt virus than other cultivars in
the USA.
Peanut stripe, both aphid-transmitted and seed-borne, is composed of
strains which can be distinguished on the basis of differential host reaction.
Over 9000 lines of A. hypogaea were screened at two sites in Indonesia
without any resistance being found (Nigam et al., 1991). A few wild species
have shown resistance with one species, A. cardenasii, being immune
(Stalker and Moss, 1987).
Peanut mottle virus (PMV) disease of groundnut is widespread and
generally present in varying intensity in all major groundnut-growing areas
of the world. It can cause up to 30% loss in yield (Kuhn and Demski,
1975). Because PMV's foliar symptoms are inconspicuous, it has not
received much attention in crop improvement programmes. Infected plants
show mild mottling and vein clearing in newly formed leaves. Older leaves
show upward curling and interveinal depression with occasional dark green
islands. Infected plants are not severely stunted and older plants seldom
show typical symptoms. The virus is sap-transmitted and its vectors are
Aphis craccivora, A. gossypii and Myzus persicae among others. It is also
seed-transmitted in a range from 0.1 % to 3.5%, depending on the ground-
nut genotype (Ghanekar, 1980).
From a 5-year study on PMV epidemiology in Georgia, USA, Kuhn and
Demski (1975) concluded that the initial inoculum of the disease in the
field came from seedlings originating from infected seeds. Taking a lead
from this observation, the groundnut group at ICRISAT adopted the
approach of combining resistance/tolerance to PMV with absence of seed
transmission in the disease resistance breeding. Limited breeding efforts
are under way to achieve this objective.
A rapid method of field inoculation has been developed (Ghanekar,
1980), by means of which about 1000 plants can be inoculated in 1 hour
with 80% infection frequency. The method involves the spraying of ex-
tracts from infected leaves, prepared in phosphate buffer containing celite
and mercaptoethanol, onto test plants through a fine nozzle under pressure
of 50 psi. More than 2500 germplasm lines of A. hypogaea have been
screened in the field. No line has shown resistance to the virus; however,
many germ plasm lines suffered much lower yield loss than controls. Two
germplasm lines, NC Ac 2240 and NC Ac 2243, have shown significantly
low yield loss due to disease over years (ICRISAT, 1983). A few breeding
lines have also shown tolerance to the disease.
Fifty wild Arachis species accessions have been screened for PMV
resistance under glasshouse conditions using mechanical leaf rub and air
brush inoculations. Of these, only two species, A. chacoensis (10602) and
A. pusilla (12911), remained free from infection even after repeated graft
inoculations (Subrahmanyam et al., 1985).
Seeds of PMV-infected plants of several germplasm lines were screened
in the laboratory for virus presence, using ELISA (Reddy, 1980). With this
technique, 1000 seeds can be screened in 2 days. A small portion of
cotyledon is adequate for the test. Two rust-resistant germplasm lines, EC
76446(292) and NC Ac 17133(RF), have failed to show any seed trans-
mission in repeated tests over years on seeds totalling more than 13 000
(ICRISAT, 1988). A recently released Indian cultivar and many breeding
lines with these rust-resistant parents in their ancestry also have shown no
seed transmission. Lines with low yield loss and no seed transmission
characteristics have been crossed and advanced generation lines are in field
tests for measuring yield loss due to the disease. Promising lines from these
tests will be studied for non-seed transmission in the laboratory.
Peanut clump virus (PCV) disease has been reported from West Africa
(Trochain, 1931; Bouhot, 1967) and India (Sundararaman, 1926; Reddy et
al., 1979). The virus is soil-borne and seed-transmitted (ICRISAT, 1986).
Infected plants are severely stunted with small, dark green leaves. The
young tetrafoliolate leaves show mosaic mottling and chlorotic rings. Roots
become dark in colour and the outer layers peel off easily. Most of the
early-infected plants fail to produce pods. Even in case of late infection,
losses of up to 60% are recorded. The virus has many serologically distinct
isolates which produce varying severity of disease on groundnut varieties
and different reactions on diagnostic hosts. A few soil fungi and nematode
species have been suspected as possible vectors of the virus. Studies in
India have shown that Polymixa graminis, a soil fungus, can transmit the
virus (ICRISAT, 1988).
The disease occurs in both warm summer and rainy season crops. The
extent of area infected with the disease is not well documented. Individual
fields can become severely infected with the virus, forcing farmers to
abandon groundnut cultivation in those fields. Chemicals such as
Nemagone®, Temik®, and Carbofuran® can greatly reduce disease and
increase yields. However, these chemicals are expensive for most farmers
of the semi-arid tropics. Solarization treatment of the infected areas of the
field greatly reduces the disease incidence (ICRISAT, 1987).
More than 7000 germplasm lines of the cultivated groundnut species
Arachis hypogaea have been screened in farmers' diseased fields in the
Indian states of Punjab and Andhra Pradesh. None of these lines showed
resistance to the virus. A few lines showed tolerance to the disease as they
did not suffer severely in growth and yield. Of 38 wild Arachis species and
their 200 interspecific derivatives tested, only Arachis species 30036 did not
become infected in the field (ICRISAT, 1985). Due to the genetic com-
plexity of virus populations and lack of high-level tolerance in germplasm,
no resistance breeding activity has been started for this disease.
14.2.3 Soil-borne pathogens

Screening of groundnut germplasm for resistance to soil-borne diseases has
been less extensive than screening for resistance to foliar fungal pathogens
because of the local prevalence of most soil-borne diseases. Nevertheless,
variation for reaction to several soil-borne diseases has been found in
groundnut. Resistance to bacterial wilt caused by Pseudomonas soiana-
cearum was reported in the 1920s by Dutch scientists working in East Java
(Indonesia) (Buddenhagen and Kelman, 1964). The disease occurs in
several Asian and African countries but significant losses are reported only
for Indonesia and China. Numerous resistant genotypes have been ident-
ified in those two countries (Nigam et ai., 1991).
From screening in North Carolina, USA, a few virginia and several
spanish genotypes were reported to be resistant to Cyiindrocladium crota-
iariae, which causes cylindrocladium black rot disease (CBR) (Green et al.,
1983). NC 3033, a line resistant to CBR, was also found to be resistant to
Sclerotium rolfsii, the causal organism of southern stem rot (Beute et ai.,
1976).
Toalson, PI 341885 and TxAG-3 (a selection from PI 365553) were
found to be resistant to southern stem rot and pythium pod rot caused by
Pythium myriotyium in Texas (Smith et ai., 1989). Resistance has also been
found to Sclerotinia minor in screening studies in Oklahoma and Virginia
(Coffelt and Porter, 1982). Sources of resistance include Chico, germplasm
from Texas (TX 498731, TX 798736, TX 804475), germplasm from
Virginia (TRC 02056-1), and seven accessions from China (PIs 467829,
476831,476834,476835,476842,467843, and 467844) (Wynne et ai., 1991).
14.2.4 Aflatoxin
Environment and cultural practices can make groundnut plants and seeds
prone to invasion by toxigenic species of Aspergillus (A. flavus and A.
parasiticus) - discussed also in Chapters 10 and 13. Seeds may be contami-
nated with aflatoxin before harvest, during post-harvest curing and drying,
or during storage. In some regions the problem develops predominantly
post-harvest while in others it is largely a preharvest phenomenon. Several
recommendations have been made regarding cultural practices, curing and
drying procedures, and storage conditions to minimize seed invasion by A.
flavus. However, these recommendations have not been widely adopted in
developing nations where groundnut production is subject to the vagaries
of the weather.
Aflatoxin contamination was considered a post-harvest problem and
received little attention in breeding programmes until it was reported by
Mixon and Rogers (1973) that two germplasm lines, PI 337409 and PI
337394F, were resistant to seed invasion and colonization by A. flavus.
Their screening method used rehydrated, sound, mature seeds inoculated
artificially with A. flavus conidia in an environment favourable to fungal
growth. They suggested that this resistance to invasion and colonization to
A. flavus, associated with the seed coat, could be an effective means of
preventing aflatoxin contamination. Varietal resistance to aflatoxin pro-
duction in groundnut seed also was reported by others (Rao and Tulpule,
1967; Kulkarni et al., 1967). These findings stimulated further research on
varietal resistance in several countries.
Resistance to A. flavus in groundnut may operate at three sites in the
plant: the pod, the seed coat and the cotyledons. Genetic variation in pod
resistance to A. flavus has been attributed to differences in pod-shell
structure (Zambetakkis et al., 1981), presence of antagonistic microflora in
the shell (Kushalappa et al., 1979; Mixon, 1980), and the presence of thick-
walled parenchyma cells (Pettit et al., 1977). Field screening for pod
resistance has been limited somewhat due to the problem of consistently
reproducing the environmental conditions required to promote infection.
Infection of seeds from the field may be assessed by surface sterilizing
seeds from mature intact pods and then incubating them under conditions
conducive to fungal growth. Disease reaction is typically expressed as the
percentage of seeds exhibiting colonization.
Seed-coat resistance has also been associated with different character-
istics such as the compact arrangement of testa cells and small hilum with
little exposure of parenchyma cells (Taber et al., 1973), waxes deposited on
the testa (LaPrade et al., 1973), 5,7-dimethoxyisoflavone (Turner et al.,
1975), tannin (Sanders and Mixon, 1978; Lansden, 1982; Karchesy and
Hemingway, 1986), and total soluble amino compounds and arabinose
content (Amaya et al., 1980). However, Jambunathan et al. (1989) did not
find significant correlation between seed colonization and polyphenol con-
tent in seed coat. Procedures for assay of in vitro seed colonization by A.
flavus (IVSCAF) utilize artificial inoculation to ensure uniform exposure of
seeds to the pathogen. Sound mature seeds from intact, dried pods are
surface sterilized, imbibed, and inoculated with a conidial suspension of a
toxigenic strain of A. flavus or A. parasiticus, then incubated to promote
mycelial growth (Mixon and Rogers, 1973; Mehan et al., 1981).
Many sources of resistance have now been reported for preharvest seed
infection, in vitro seed colonization and aflatoxin production (Table 14.5).
These include PI 337409, PI 337394F, UF 71513, J 11, Ah 7223, U-4-47-7,
55-437, and 73-30 for preharvest field infection and colonization and
aflatoxin production. J 11 is grown commercially in India, as are 55-437 and
73-70 in Senegal and other West African nations. Three lines with resist-
ance to IVSCAF (PI 337394F, PI 337409 and J 11) have been evaluated in
more than one country. J 11 exhibited resistance to seed infection in India
and the USA. PI 337409 was resistant in tests in Senegal and India, but was
susceptible in the USA (Kisyombe et al., 1985). Mixon (1976) recorded
percentage colonization of seeds in the Fl and F2 generations of crosses
between PI 337409 and PI 331326, a susceptible line. Broad-sense heritabi-
lity was estimated at 78.5%. Based on diallel and factorial matings con-
ducted at ICRISAT Center, Vasudeva Rao et al. (1989) reported that UF
71513, Ah 7223, PI 337394F and PI 337409 had good combining abilities
for seed-coat resistance. Resistance to IVSCAF in breeding lines
TABLE 14.5 Sources of resistance to Aspergillus flavus or A. parasiticus
Source of Type of Country

resistance resistance where used Reference
1-4 IVSCAF India Ghewande et al., 1989

1-7 IVSCAF India Ghewande et al., 1989
55-437 Field infection Senegal Waliyar and Bockelee-Morvan,
1989
Zambetakkis et al., 1981
IVSCAF Senegal Zambetakkis et al., 1981
73-30 Field infection Senegal Waliyar and Bockelee-Morvan,
1989
Zambetakkis et al., 1981
IVSCAF Senegal Zambetakkis et at., 1981
73-33 Field infection Senegal Waliyar and Bockelce-Morvan,
1989
Zambetakkis et at., 1981
IVSCAF Senegal Zambetakkis et al., 1981
A. cardenasii Aflatoxin India Ghewande et al., 1989
production
IVSCAF India Ghewande et al., 1989
A. duranensis Aflatoxin India Ghewande et al., 1989
production
IVSCAF India Ghewande et al., 1989
Acc 63 IVSCAF Philippines Pua and MedalJa, 1986'
Ah 6487 IVSCAF China Tsai and Yeh, 1985
Ah 7223 Field infection India Mehan et at., 1986b, 1987
IVSCAF India Mehan and McDonald, 1980
Ghewande et al., 1989
AR-1 IVSCAF USA Mixon, 1983b
Basse IVSCAF China Tsai and Yeh, 1985
C 116(R) IVSCAF China Tsai and Yeh, 1985
C 184 IVSCAF China Tsai and Yeh, 1985
Celebes IVSCAF Philippines Pua and Medalla, 1986
CES 48-30 IVSCAF Philippines Pua and Medalla, 1986
CGC7 IVSCAF India Ghewande et al., 1989
CGC-2 IVSCAF India Ghewande et al., 1989
Darou IV Pod infection Senegal Zambetakkis, 1975
F-7 IVSCAF China Tsai and Yeh, 1985
Faizpur IVSCAF India Mehan and McDonald, 1980
GE 652 IVSCAF China Tsai and Yeh, 1985
GFA-1 IVSCAF USA Mixon, 1983a
GFA-2 IVSCAF USA Mixon, 1983a
Jll Field infection India Mehan et at., 1986b, 1987
Ghewande et al., 1989
USA Kisyombe et al., 1985
TABLE 14.5 Cant.
Source of Type of Country

resistance resistance where used Reference
M395 IVSCAF China Tsai and Yeh, 1985

Maria-B IVSCAF China Tsai and Yeh, 1985
Monir 240-30 IVSCAF India Mehan and McDonald, 1980
NC449 IVSCAF China Tsai and Yeh, 1985
NC482 IVSCAF China Tsai and Yeh, 1985
PI 196621 IVSCAF China Tsai and Yeh, 1985
PI 196626 IVSCAF China Tsai and Yeh, 1985
PI337394F Field infection India Mehan et aI., 1986b, 1987
Senegal Waliyar and Bockelee-Morvan,
1989
Zambetakkis et at., 1981
Senegal Zambetakkis et at., 1981
USA Mixon and Rogers, 1973
PI 337409 Field infection Senegal Zambetakkis, et at., 1981
Senegal Zambetakkis et at., 1981
USA Kisyombe et at., 1985
Mixon and Rogers, 1973
PI 339407 Field infection India Mehan et at., 1986b, 1987
Senegal Waliyar and Bockelee-Morvan,
1989
RMP 12 IVSCAF China Tsai and Yeh, 1985
Roxo (Sal) IVSCAF China Tsai and Yeh, 1985
S 230 IVSCAF India Ghewande et at., 1989
Shulamith Pod infection Senegal Zambetakkis, 1975
Sp.218 IVSCAF China Tsai and Yeh. 1985
Sp.424 IVSCAF China Tsai and Yeh, 1985
U4-47-7 Field infection India Mehan et at., 1986b, 1987
U4-7-5 Aflatoxin India Mehan et at., 1986a
production
UF 71513 Field infection India Mehan et at., 1986b, 1987
IVSCAF USA Bart et at., 1978
UPLPn4 IVSCAF Philippines Pua and Medalla, 1986
Var 27 IVSCAF India Mehan and McDonald, 1980
VRR 245 Aflatoxin India Mehan et at., 1986a
production
developed in India has remained stable over years and locations (Vasudeva
Rao et al., 1989).
In the United States, there is controversy as to the value of IVSCAF in
practical control of Aspergillus contamination. Wilson et al. (1977) found
production of aflatoxin in PI 339396F and PI 339407 to be similar to
IVSCAF-susceptible genotypes PI 334360 and Florunner when seed lots
were stored under high humidity. All lots exhibited 2-3% infection of
seeds by Aspergillus spp. prior to storage. None of the lots was inoculated.
Davidson et al. (1983) compared aflatoxin contamination in farm-grown
samples of Florunner with Sunbelt Runner, a cultivar selected for resist-
ance to IVSCAF. Seeds of Sunbelt Runner sampled prior to storage
exhibited levels of natural infection and aflatoxin production comparable
to Florunner. Seed-coat resistance is operative only in seeds with intact
testae. The conditional nature of this resistance limits its utility under field
conditions. Its effectiveness is reduced by mechanical operations causing
pod and seed damage or by faulty curing, drying and storage conditions.
Genetic variation has been observed for the ability of groundnut cotyle-
dons to support production of aflatoxins (Rao and Tulpule, 1967; Kulkarni
et al., 1967; Doupnik, 1969; Aujla et al., 1978; Doupnik and Bell, 1969;
Nagrajan and Bhat, 1973; Tulpule et al., 1977). Very little is known about
the mechanism of resistance to aflatoxin production. Several studies have
reported effects of fungal nutrition on toxigenesis by Aspergillus spp.
grown on defined media. Payne and Hagler (1983) observed differences in
the growth of Aspergillus spp. on media containing different amino acids.
Casein, proline, asparagine and ammonium sulphate supported fungal
growth and toxin production better than did tryptophan or methionine.
Venkitasubramanian (1977) found toxin production to be enhanced on
defined media containing casamino acids rather than urea or ammonium
nitrate as the nitrogen source. Maggon et al. (1973) studied the effects of
micronutrients on aflatoxin biosynthesis, finding that toxin production was
stimulated by copper but inhibited by cadmium, barium and vanadium.
Screening methods for aflatoxin production are similar to those used for
seed colonization. Some researchers have removed the testa of the seed
prior to inoculation in order to remove any barrier to infection contained
therein. Inoculated seeds are incubated and aflatoxin measured using thin
layer chromatography (Mehan and McDonald, 1980). Mehan et al. (1986)
identified U4-7-5 and VRR 245 as resistant to production of aflatoxin.
U -4-7 -5 and VRR 245 do not support high levels of aflatoxin production
but are susceptible to colonization and seed invasion. A previous report of
two wild species, A. cardenasii and A. duranensis, supporting production
of only trace levels of aflatoxin (Ghewande et al., 1989) was not confirmed
in subsequent screening performed at ICRISAT (Mehan et al., 1992).
A. flavus is a weak pathogen. Its ability to invade intact pods and seeds is
strongly influenced by environmental conditions during pod and seed
development. Developing pods must be predisposed to infection by the
occurrence of water stress in the soil surrounding them and by high soil
temperatures (38-40 DC) in the podding zone (Cole et af., 1989). These
conditions weaken the host plant and suppress the growth of soil microbes
antagonistic to or competitive with A. fiavus. At ICRISAT Center, field
screening for resistance to preharvest infection is conducted in the post-
rainy season; severe drought stress is imposed by withholding irrigation
late in the growth cycle.
14.2.5 Insect pests

Groundnut is subject to reduction of yield and quality due to feeding by
insects and arachnids on leaves, pegs, pods and seeds. In addition to
causing damage directly, some insects serve as vectors of viral diseases.
Insects of global importance include aphids, thrips, jassids and Spodoptera.
Leaf miner, Hilda, Helicoverpa and other lepidopterous species present
problems in specific regions. In Asia and Africa, white grub is the most
economically important pod-feeding pest, but termites, millipedes and ants
may also damage pods in specific regions. In the USA, lesser cornstalk
borer (Efasmopafpus fignosellus) , white-fringed beetle (Graphognathus
spp.) and southern corn rootworm (SCR, Diabrotica undecimpunctata
howardii) are the primary agents of damage to pegs and pods. Damage
from pod feeders not only reduces yield but also permits entry into the pod
of soil-borne pathogens such as A. fiavus.
Sources of resistance to most insect pests have been identified (Lynch,
1990; Wightman et al., 1990; Nigam et af., 1991) although levels of resist-
ance do not approach immunity. Some sources exhibit resistance to more
than one pest:
NC6
GP-NC 343
NC Ac 01705
NC Ac 02142
NC Ac 02214
NC Ac 02230
NC Ac 02232
NC Ac 02240
NC Ac 02242
NC Ac 02243
NC Ac 02460.
These sources of resistance trace ancestry to PI 121067 or to X-irradiated
leaf mutants of NC 4 selected by W.e. Gregory and D.A. Emery at North
Carolina State University in the 1950s. Several have dense, elongated or
erect trichomes on leaflet surfaces.
Dwivedi et af. (1986) reported predominantly non-additive genetic vari-
ance for trichome characters. Additive genetic effects were important for
trichome length and jassid damage. Holley et al. (1985) found additive
genetic effects to predominate for resistance to a complex of insect pests
(thrips, jassids and Helicoverpa) in North Carolina. Several breeding lines
and cultivars resistant to foliar diseases (ICG [FDRS] 4, ICG [FDRS] 10,
ICGV 86590, GP-NC 343 and NC 6) also exhibit tolerance to one or more
insect species such as Spodoptera, leaf miner or jassids.
14.3 BREEDING METHODS
Early breeders of groundnut used mass selection to exploit natural vari-

ation in local cultivars. This method was commonly used in the USA
during the 1950s but was gradually replaced by use of mutagenesis or
hybridization as means of creating new genetic variation. It is interesting to
note that mass selection is still used to some extent today, especially in
conjunction with genetic stocks introduced from outside the USA. It is
common for groundnut cultivars to exhibit some phenotypic variation in
the field. This could be the result of segregation within the progeny of the
last single plant selected in the course of cultivar development, of segre-
gation and assortment following natural hybridization between pure-line
components of a genetically heterogeneous but phenotypically homogene-
ous cultivar, or of duplication or deletion of chromosomal segments follow-
ing the occasional formation of quadrivalents in the first meiotic division of
the tetraploid groundnut. The most recently released American cultivar
developed by mass selection was Avoca 11, a virginia cultivar selected from
Florigiant and released in 1976.
The method most commonly used by groundnut breeders is the pedigree
method. This allows the breeder to practise selection for highly heritable
characters such as pod and seed size and shape, plant type and testa colour
in early segregating generations. Because these traits determine market
type and conformation to local standards, they are generally the focus of
intensive early-generation selection. This practice serves to reduce quickly
the size of individual segregating populations. Only when the desirable
plant, pod and seed type have been recovered is emphasis placed on
quantitative characters such as yield and seed composition.
Modified pedigree (single-seed descent) procedures and recurrent selec-
tion have been used in groundnut (Wynne, 1975; Hildebrand, 1985) but
are not the methods of choice. Despite the recommendation of Brim
(1966) that single-seed descent be used to allow segregating populations to
resolve into collections of pure lines before selecting even for qualitative
traits, groundnut breeders have continued to favour the pedigree method.
The basis for this preference may lie in the space-intensive nature of plot
work in groundnut. In modified pedigree procedures, the breeder must for
several generations carry forward large populations of plants, selecting a
single pod from each at random. This necessitates planting at population
Regional progress 575
densities sufficiently low to allow identification of individual plants.
Particularly in populations segregating for spreading growth habit, indi-
vidual ground nut plants may occupy large areas relative to small grains or
grain legumes bearing aerial fruit.
Backcross breeding has not been used extensively in groundnut due to
the paucity of simply inherited disease and insect resistances. This method-
ology may find greater favour in the future as recently identified resistances
to rust and late leaf spot or characters such as the Florida high OIL ratio
are transferred into existing cultivars that meet exacting standards of
processors and consumers. Backcrossing augmented by use of molecular
techniques for identifying heterozygotes may be used for transfer of genes
introduced into A. hypogaea through transformation procedures, although
it is to be hoped that transformation protocols insensitive to the recipient
genotype will be developed, allowing independent transformation of any
existing cultivar.
Development of genetic maps utilizing allozymes, RFLPs or random
amplified polymorphic DNAs (RAPDs) as markers has promised to re-
solve the poly- or oligogenically inherited, quantitative traits such as yield
to essentially qualitative traits by allowing the breeder to identify chromo-
somal segments bearing genes with measurable effects on the quantitative
traits. Such methodology requires the genomic map to be saturated with
markers, i.e. that there be markers exhibiting polymorphism in the segre-
gating population of interest at average intervals of 5-20 centimorgans.
Unfortunately, cultivated groundnut exhibits very little polymorphism for
allozymes or RFLPs, making this approach to groundnut improvement
impractical at present. On the other hand, the diploid wild species of
Arachis exhibit large amounts of polymorphism for allozymes and RFLPs.
It may be possible to utilize these markers for construction of a genomic
map in the diploid species and to monitor the incorporation of wild species'
germplasm in populations of cultivated groundnut. The foremost potential
obstacle to use of molecular markers in wild species is the possibility of
abnormal recombination between homologous chromosomes of related
species, especially if the specific genomes are differentiated by structural
rearrangements such as inversions or translocations.
14.4 REGIONAL PROGRESS
14.4.1 Africa
An important cash and food crop in Africa, ground nuts have declined
there in terms of area, yield and productivity over the past 20 years. Two
epiphytotics of groundnut rosette virus in West Africa in 1975 and 1987
almost wiped out the crop in many countries, leaving not even enough seed
for farmers to plant their next crop. The changing rainfall pattern in West
Africa and other parts of the continent has resulted in reduction of the
length of the rainy season and forced groundnut out of cultivation in
desiccated areas where it once was a major crop.
Groundnut research in Africa began during the colonial period. Colonial
governments made serious efforts to establish and increase groundnut
production in their colonies to meet the increasing demands of home
industries and population. This effort was strengthened during and after
World War II, when shortages in Europe became acute. During that
period, much research was conducted in Burkina Faso, Senegal, Nigeria,
Uganda, Tanzania, Zambia, Zimbabwe, Malawi, Sudan and Zaire. After
the decolonization of Africa the same impetus in research could not be
maintained by newly independent nations. Civil strife, lack of physical
resources, deteriorating infrastructure and lack of trained scientists and
technicians resulted in the near-death of many national research pro-
grammes. Work was discontinued, valuable germplasm lost, records des-
troyed and cultivars mixed.
Over the past two decades many national programmes have been revita-
lized with the support of international organizations and donor agencies
such as FAO, UNDP, ICRISAT, ODA, USAID, IDRC, IRHO, IRAT,
GTZ and others. However, the revival process has been slow and many
national programmes collapse as soon as financial support by donor agen-
cies is withdrawn. Many countries have better trained and qualified scien-
tists, but the lack of the resources necessary to conduct needed research
continues to plague many national programmes. Lately the World Bank
has taken interest in restructuring the national agricultural apparatus in
Africa. IRHO, IRAT, and ICRISAT through its regional programmes in
Malawi and Niger have made long-term commitments to the region and are
making efforts to strengthen national programmes. USAID's Peanut
Collaborative Research Support Program (CRSP) has been involved in
development of West African peanut research for the past 10 years (Peanut
CRSP, 1990).
Most results of research conducted in Africa are confined to annual
reports of individual projects. Very little is published in international
journals. Due to poorly developed seed production, distribution and exten-
sion programmes, most new cultivars and new cultural practices have not
been adopted by producers at large. Uncertain tenure of land, lack of
price support and unavailability of credit have discouraged farmers from
increasing their groundnut production. Importing nations, particularly the
European Community, have established extremely low tolerances for afla-
toxin in imported groundnuts - levels difficult to meet for developing
nations with generally poor storage and handling facilities. Export markets
for African countries have declined due to poor quality and irregular
supply of groundnuts.
From reports of 30 African nations published in proceedings of work-
shops conducted by ICRISAT and in reports of other organizations, the
most important constraints on increased groundnut production in Africa
(excluding socioeconomic factors) include important biotic stresses such as
foliar fungal diseases (early leaf spot, late leaf spot, rust), viral diseases
(groundnut rosette virus, peanut clump virus and peanut mottle virus),
arthropod pests (aphids, thrips, leaf miner, Spodoptera, jassids, white
grubs, Hilda patruelis, termites and millipedes) and other animal pests
(nematodes, rats, squirrels and monkeys).
Abiotic stresses of primary importance are drought and poor soil fer-
tility. Other stresses are restricted in distribution to one or two countries.
They include bacterial wilt in Uganda; Aleetra species; phanerogamic root
parasitic weeds in Nigeria and Malawi; acid soils in Zaire, Zambia and
Malawi; and Phoma araehidicola in Zimbabwe.
Breeding objectives of the national programmes in Africa can be sum-
marized as development of high-yielding oil type and/or confectionery
cultivars with adaptation to specific agroecological conditions and resist-
ance to the stresses constraining yield. Resistance to leaf spots, rust, A.
flavus, ground nut rosette virus, tolerance to drought and early maturity
rate high in most breeding programmes. Very little effort has been
expended on breeding for resistance to animal pests.
Breeding methods employed in Africa are similar to those used else-
where in the world. Programmes with limited resources or technical ex-
pertise for hybridization and selection rely primarily on introduction and
pure-line selection within local landraces. International institutes such as
ICRISAT and bilateral programmes such as IRHO and USAID's Peanut
CRSP continue to be major sources of new genetic material in African
national programmes.
Hybridization has been used in only a few national programmes and only
intermittently in those. Countries with stronger programmes distribute
their cultivars to neighbouring nations and to nations sharing common
linguistic or economic ties with a former colonial power. For example,
Burkina Faso and Senegal have shared their cultivars with other countries
in francophone West Africa while Zambia has provided cultivars to nations
of southern Africa with ties to the UK. In programmes using hybridization,
pedigree selection has been the most commonly used method of generation
advance. The backcross method has been used in breeding for disease
resistance. Zimbabwe's national programme used a modified pedigree
method (single-seed descent) to develop two cultivars (Hildebrand, 1985).
The Zambian national programme has also used single-seed descent.
Interspecific hybrids obtained from the University of Reading, North
Carolina State University and ICRISAT have been evaluated for resist-
ance to foliar diseases in Malawi and Zimbabwe and for resistance to foliar
diseases and insect pests in Nigeria. A programme of mutation breeding
was initiated in Uganda to create variability for selection because the
breeder there found the time required for emasculation and pollination to
be excessive (Busolo-Bulafu, 1990).
Increased desertification in sub-Saharan Africa has made breeding for
drought resistance a primary objective in that region. The Senegalese
programme at the Bambey centre of the Institut Senegalais de Recherches
Agricoles (ISRA) has developed many cultivars with improved resistance
to drought, including 47-16, 50-127, 73-33 and 55-437 (Bockelee-Morvan et
ai., 1974). Adaptation to dry climate was achieved by shortening the
growing cycle of the breeding lines using 'Chico' as a source of early
maturity and screening lines for tolerance to drought (Gautreau and De
Pins, 1980). Recently, a joint programme between ISRA and the Sebele
Research Station of the Botswana Department of Agriculture at Gaborone
was initiated to improve drought tolerance in groundnut. Two crops are
grown each year, one in Senegal and one in Botswana. Eight cultivars
(virginia types 47-16,57-422,59-127 and 73-33 and spanish types 49-20,55-
437, 68-111 and TS 32-1) were used as parents in a convergent (pyramidal)
crossing scheme (Mayeux, 1987). Drought-tolerant germplasm developed
at ICRISAT Center near Hyderabad, India, has been introduced into
southern and West Africa.
Breeding for resistance to rust and late leaf spot is ongoing in many
national programmes including Burkina Faso, Malawi, Nigeria, Senegal,
Zambia and Zimbabwe. These continue to emphasize the introduction of
improved resistant germplasm from ICRISA T and the USA. 'RMP 91', a
GRV-resistant cultivar developed in Burkina Faso, was found to be toler-
ant to leaf spots. A few programmes have crossed introduced sources of
resistance with local cultivars. No cultivar with resistance to foliar fungal
pathogens has been released in Africa to date.
African cultivars have been screened to identify resistance to early leaf
spot, but no resistant cultivars have been found. At the SADCC-ICRISAT
Regional Groundnut Program in Malawi, several germplasm lines and
advanced breeding lines have been found to retain foliage longer than
checks under intense disease pressure (Bock, 1987). These sources of
resistance to defoliation are being intermated to improve the level of
resistance. Of the Arachis species screened in Malawi, A. sp. 30003
exhibited a high level of resistance to early leaf spot. Unfortunately, this
diploid species cannot be crossed directly with A. hypogaea.
Breeding for resistance to groundnut rosette virus has been remarkably
successful in Africa. Resistance to GRV was identified in local landrace
cultivars in Burkina Faso by de Berchoux (1958) ,who later (1960) showed
that the resistance was governed by two independent recessive genes. The
resistance operates equally against both chlorotic (de Berchoux, 1960) and
green (Harkness, 1977) rosette. Nigam and Bock (1990) confirmed de
Berchoux's observations and described an effective field screening tech-
nique for rosette. Utilizing resistance from landraces, IRHO breeding
programmes in Burkina Faso and Senegal have developed several GRV-
resistant cultivars including RMP 12, RMP 91, 69-101, KH-149A and
KH-241D. The last two cultivars are spanish type; the others are virginia
type. In southern Africa, the Malawi national programme developed a
GRV-resistant cultivar, RGl. For many regions in Africa, current empha-
sis in breeding for resistance to rosette is on transferring resistance into
early-maturing cultivars. The SADCC-ICRISA T Regional Groundnut
Program and the Nigerian national programme are actively involved in
GRV-resistance breeding.
Other than locallandraces, the genetic source that has contributed most
to varietal development in Africa is Mani Pintar. The history of this line
illustrates the powerful role of introduction in crop improvement. Mani
Pintar was collected from a market place in La Paz, Bolivia, by Stephens
and Hartley in 1947 (Hartley, 1949). The name is undoubtedly a corruption
of 'manl pintado' or 'painted groundnut'. The characteristic features of the
line are red-and-white variegated testa and spreading bunch growth habit
(cultivar group N ambyquarae). The original seed sample was shared by the
Queensland Department of Agriculture and Stock in Australia and the
USDA. In the USA, the accession was assigned plant introduction number
PI 162404. In 1955 the accession was introduced to the Mount Makulu
Research Station in Zambia, where pure line selection was practised in
subsequent years. A single-plant selection with solid red testae led to the
release of 'Makulu Red' in 1961 (Smartt, 1978). Mani Pintar and Makulu
Red were introduced into Zimbabwe in 1960. Sigaro Pink, a variant with
pink testae, arose from Makulu Red, presumably as a result of natural
hybridization, and was released in Zimbabwe in 1968-69. Further selection
within Sigaro Pink gave rise to Apollo in 1972-73 and Egret in 1975. Mani
Pintar is also one of the parents of GRV-resistant cultivars RMP 12 and
RMP 91, which are very popular in West Africa.
There are more than 65 released cultivars reported in the literature from
Africa. However, only a few are grown on a large scale and are pan-
African in nature (Table 14.6). Most of the common cultivars of West
Africa were developed by ISRA's Centre Nationale pour les Recherches
Agricoles (CNRA) at Bambey, Senegal, and by IRHO in Burkina Faso.
14.4.2 East Asia

China, Japan and South Korea are the major groundnut-growing countries
in east Asia. China is the leading groundnut producer in the world. In
1989, the groundnut area in the country was 2946000 ha and the total
production was 5 362 000 t with an average yield of 1815 kg/ha. Compared
with the 1970s, the groundnut area in China in the 1980s increased by 50%,
the production by 124% and the yield by 48%. In this period, old cultivars
were replaced by new improved cultivars in 95% of the groundnut area of
the country. Groundnut cultivation in China is concentrated in the north-
ern region, which accounts for 60% of the total groundnut area. Shan dong
Province in the northern region is the leading groundnut-producing pro-
vince in China with an average pod yield of 2.7 t/ha. Other important areas
are the southern (21 %) and central (12%) region.
TABLE 14.6 Cultivars released in Africa
Cultivar Type Origin or pedigree Year Description

Burkina Faso (IRHO. Niangoloko Station)
Te.3 Spanish Selection from a local 1958 90-day cycle, erect growth habit, medium leaflet size, semi-compact fruiting habit, small
population from Upper (70-80 g/lOO) 2-seeded pods with crest and slight constriction, no beak, small (38-40 g/100)
Volta flattened seeds, salmon pink testa, 67-70% meat content, 47-48% oil content. no seed
dormancy. 41-43% oleic acid content, 33-35% linoleic acid content, resistant to drought.
Used in Benin, Burkina Faso.
RMP 12 Virginia F9 selection following 1963 135-150-day cycle, semi-spreading growth habit, medium leaflet size, compact fruiting habit,
hybridization, 1036 / medium (80-90 g/100) 2-seeded pods with no crest or constriction, marked reticulation,
Mani Pintar moderate beak, medium (50-55 g/lOO) oblong flattened seeds, pink variegated testa. 72%
meat content. 49% oil content, 98% seed dormancy, 55-58% oleic acid content, 24-26%
linoleic acid content, sensitive to drought, excellent resistance to GRV. susceptible to rust.
Used in Benin, Burkina Faso, Mozambique, Nigeria.
RMP91 Virginia F9 selection following 1963 135-150-day cycle, semi-spreading growth habit, medium leaflet size, compact fruiting habit,
hybridization, 48-37 / small (75-85 g/100) 2-seeded pods with no crest or constriction, marked reticulation, moderate
Mani Pintar beak, small (48-50 g/100) oblong seeds, pink testa, 68% meat content, 48% oil content, 98%
seed dormancy, 55-58% oleic acid content, 24-26% linoleic acid content, sensitive to drought,
excellent resistance to GRV, tolerant to early and late leaf spots.
Used in Benin, Burkina Faso, Cameroon, Nigeria.
KH-149 A Spanish F7 selection following 1964 90-day cycle. semi-spreading growth habit, medium leaflet size, semi-compact fruiting habit,
hybridization, GH 119- small (65-75 g/100) 2-seeded pods with deep constriction, no crest. slight beak, small (30-
7.1II-III / 91 Saria 35 g/lOO) oblong seeds, red testa, 67-70% meat content. 48-50% oil content, no seed
dormancy, 37-39% oleic acid content, 34-36% linoleic acid content, low resistance to
drought. resistant to GRV.
Used in Benin, Burkina Faso, Niger.
KH-214 D Spanish F7 selection following 1964 90-day cycle, semi-spreading growth habit. medium leaflet size, semi-compact fruiting habit,
hybridization, GH medium (80-90 g/lOO) 2-seeded pods with very slight constriction. no crest, moderate beak,
1185.2 III 91 Saria small (35-40 g/lOO) flattened seeds, red testa, 70% meat content, 49-50% oil content, no seed
dormancy. 38-40% oleic acid content. 35-37% linoleic acid content, resistant to drought,
resistant to GRV.
Used in Benin, Burkina Faso.
TABLE 14.6 Cant.

TS 32-1 Spanish Selection following 1966 90-day cycle, erect growth habit, medium leaflet size, semi-compact fruiting habit, small
hybridization, Spanlex (70-80 g/100) 2-seeded pods with moderate constriction, no crest, slight beak, small (35-38 gI
Te. 3 100) slightly flattened seeds, pink testa, 68-70% meat content, 50-51% oil content, no seed
dormancy, 44-46% oleic acid content, 31-33% linoleic acid content, resistant to drought.
Used in Benin, Burkina Faso, Chad, Niger.
Congo (IRHO, Loudima)
A-124 B Valencia Selection from a local 1956 Long Manyema group, 90-day cycle, erect growth habit, large leaflet size, large (165 gIlOO)
population, Loudima 3- or 4-seeded pods with deep dorsal constriction, no crest, marked reticulation, prominent
Red beak, small (42 g/100) oblong seeds, red to purplish-blue testa, 69% meat content, 48-50% oil
content, no seed dormancy, 45-47% oleic acid content, 31-33% linoleic acid content, low
resistance to drought.
Malawi
RG 1 Virginia Selection following 1976 Resistant to GRV.
(bunch) hybridization, Makulu
Red / 48-34
Chitembana Virginia Selection following 1980 Confectionery type.

(runner) hybridization,
Chalimbana / RJ5
Mawanga Virginia Introduced from Bolivia 1980 Oil type.

(bunch)
ICGM 42 Virginia Selection following 1990 Red testa.

(bunch) hybridization,
USA 20 / TMV 10
Senegal (IRHQ)
756A Virginia Selection from a local 1951 125-day cycle, erect growth habit, medium leaflet size, semi-compact fruiting habit, large
population from the (160-200 g/100) 2-seeded pods with no constriction or crest, no beak, medium (65-75 g/100)
Casamance region of round distinctly flattened seeds, pink testa, 70% meat content, 48% oil content, complete
Senegal seed dormancy, 55-58% oleic acid content, 18-20% linoleic acid content, sensitive to drought.
TABLE 14.6 Cant.

Senegal (ISRA, Bambey CNRA)
28-206 Virginia Selection from a 1928 Samaru group. 120-day cycle. erect growth habit. medium leaflet size. compact fruiting habit.
population from medium (100-125 g/lOO) 2-seeded pods with very slight constriction. no crest. fine reticulation.
Bamako, Mali no beak. small (45-49 g/lOO) round distinctly flattened seeds. pink testa, 73% meat content.
50% oil content. complete seed dormancy. 65-68% oleic acid content. 15-18% linoleic acid
content. sensitive to drought.
Used in Cameroon. Gambia. Mali. Niger. Senegal.
47-10 Spanish Selection from a 1947 Manyema group. 90-day cycle, erect growth habit, large leaflet size. medium (105 gIlOO) 2-
population received from seeded pods with moderate dorsal constriction, prominent crest. prominent reticulation. very
Madagascar, Ambata B / prominent beak. small (45 g/100) slightly flattened seeds, salmon pink testa. 71 % meat
Morovoay content. 48% oil content, no seed dormancy, 43-45% oleic acid content. 32-33% linoleic acid
content, moderate resistance to drought. low resistance to Pythium myriotylum.
Used in Mali.
55-437 Spanish Selection from a 1955 Natal Barberton group. 90-day cycle. erect growth habit. large leaflet size. compact fruiting
population of probable habit. small (85-95 g/IOO) 2-seeded pods with slight constriction, prominent reticulation.
South American received almost no beak, small (35-38 g/100) slightly flattened seeds. pale pink testa. 75% meat
from Hungary content. 49% oil content. 30% seed dormancy. 46-49% oleic acid content. 27-30% linoleic
acid content, resistant to drought.
Used in Botswana, Cameroon. Chad, Gambia, Mali, Niger. Nigeria. Senegal. Uganda.
57-422 Virginia Selection from a hybrid 1957 105-lIO-day cycle, erect growth habit. large leaflet size. large (165-175 g/IOO) 2-seeded pods
population imported with very deep constriction, no crest, very slight reticulation prominent beak. medium (65-
from Tifton. Georgia, 69 g/IOO) slightly flattened oblong bumpy seeds, yellowish pink testa. 78% meal content. 50%
USA, F334-3-404 oil content. 95-100% seed dormancy. 50-53% oleic acid content, 27-30% linoleic acid
content, moderate resistance to drought, susceptible to late leaf spot and A. niger, tolerant to
pev.
Used in Mozambique, Niger, Senegal.
57-313 Virginia Selection from a 1957 Samaru group, 125-day cycle, erect growth habit, medium leaflet size, diffuse fruiting habit,
population from medium (125-130 g/IOO) 2-seeded pods with slight constriction. no crest. fine reticulation, no
Ouagadougou, Burkina beak, small (48-52 g/IOO) round distinctly flattened seeds. pink testa, 75% meat content, 50%
Faso oil content, complete seed dormancy, 64-67% oleic acid content, 14-17% linoleic acid
content, sensitive to drought.
TABLE 14.6 Cant.

GH 119-20 Virginia Introduced from Tifton, 1960 Jumbo group, 110-day cycle, erect growth habit, large leaflet size, fair fruiting habit, large
Georgia, USA, in 1960, (230-240 g/100) 2-seeded pods with moderate constriction, no crest, marked reticulation, large
F-l selection from (85-90 g/100) oblong seeds, pink testa, 70% meat content, 43-46% oil content, medium seed
Southeastern Runner 1 dormancy, 63-66% oleic acid content, 14-17% linoleic acid content, sensitive to drought.
Dixie Giant, 210--41/ Used in Ethiopia, Senegal.
Virginia Runner
69-101 Virginia BC,F, selection 1969 Saloum group, 125-day cycle, erect growth habit, medium leaflet size, compact fruiting habit,
follo\\'ing hybridization, medium (130 g/100) 2-seeded pods with very slight constriction, no crest, fine reticulation, no
55-455 14*28-206 beak, small (46-50 g/100) round distinctly flattened seeds, pink testa, 73% meat content, 50%
oil content, complete seed dormancy, 65-68% oleic acid content, 14-17% linoleic acid
content, sensitive to drought, resistant to GRV,
Used in Benin, Burkina Faso, Senegal.
73-27 Virginia Fs selection following 1972 Jumbo group, l20-l25-day cycle, erect growth habit, large leaflet size, fair fruiting habit, large
hybridization, 756A 1 (200, 210 g/100) 2-seeded pods with moderate constriction, no crest, slight reticulation, no
GH 119-20, Line 252 beak, large (85-90 g/100) oblong seeds, salmon pink testa, 71% meat content, good seed
dormancy, 58-61 % oleic acid content, 20-22% linoleic acid content, sensitive to drought,
used for confectionery purposes.
73-28 Virginia Fs selection following 1972 Jumbo group, l20--l25-day cycle, erect growth habit, large leaflet size, fair fruiting habit, large
hybridization, 756A 1 (190 -200 g/100) 2-seeded pods with moderate constriction, no crest, slight reticulation, no
GH 119-20, Line 255 beak, large (85-90 g/100) oblong seeds, salmon pink testa, 72% meat content, good seed
dormancy, 55-58% oleic acid content, 21-23% linoleic acid content, sensitive to drought,
used for confectionery purposes,
73-30 Spanish Fs selection following 1973 95-day cycle, erect growth habit, medium to large leaflet size, compact fruiting habit, medium
hybridization, 61-24 (100 g/100) 2-seeded pods with slight constriction, no crest, slight reticulation, no beak, small
(spanish) 159-127 (40 g/100) oblong seeds, salmon pink testa, 73% meat content, 48% oil content, complete
(virginia type Saloum) seed dormancy, 60-63% oleic acid content 18-21% linoleic acid content, resistant to drought.
73-33 Virginia F 12 selection following 1973 Fung group, 105-110-day cycle, very erect growth habit, medium leaflet size, compact fruiting
hybridization, 58-6501 habit, medium (120-125 g/100) 2-seeded pods with deep constriction, no crest, marked
59--46 reticulation, medium beak, small (50-52 g/100) oblong seeds, pink, 73% meat content, 50%
oil content, 95% seed dormancy, 58-61 % oleic acid content, 20-22% linoleic acid content,
resistant to drought.
Used in Gambia, Senegal.
TABLE 14.6 Cont.

South Africa
Natal Spanish Erect growth habit. large leaflet size. 2 seeds per pod. no constriction or beak. pale tan testa.
Common Used in Mozambique. South Africa. Tanzania. Zambia Zaire.
Tanzania
Nyota Spanish Introduced from USA 1983
in 1978 (Spancross)
lohari Spanish Introduced from India 1985

in 1980 (Robut 33-1)
Zaire
A65 Valencia Introduced from 1958 90-clay cycle. erect growth habit. rose-tan testa.
Brazil Used in Burundi. Zaire.
G 17 Valencia Selection from a local 1975 105-day cycle. erect growth habit. rose-tan testa.
landrace following
apparent natural
hybridization
Zambia (Mount Makulu Research Station)
Mani Pintar Virginia Collection from a 1955 130-140-day cycle. spreading bunch growth habit. dark green leaves. large 2-seeded pods with
market in La Paz. no constriction. pronounced beak. medium large flattened seeds. red and white variegated
Bolivia. introduced to testa.
Mt. Makulu Station in Used in Malawi. Uganda. Zambia.
1955
Makulu Red Virginia Selection from Mani 1961 130-140-day cycle. spreading bunch growth habit. dark green leaves. large leaflet size. large 2-
Pintar seeded pods with no constriction. pronounced beak. medium large flattened seeds. red testa.
67% meat content. 45% oil content. field resistance to early leaf spot.
Used in Tanzania. Uganda. Zambia. Zimbabwe.
TABLE 14.6 Cont.

Chitembana Virginia Selection from a local 1964 140-150-day cycle, runner growth habit, large leaflet size, thick stems, large coarse 2-seeded
population from pods with slight constriction, no beak, flattened seeds, dark tan testa, used for confectionery
eastern Zambia purposes.
Used in Malawi, Zambia.
Comet Spanish Introduced from USA 1984

(Comet)
MGS2 Virginia Introduced from India 1988

(runner) (M13)
Zimbabwe
Egret Virginia Selection from 1974 130-140-day cycle, spreading bunch growth habit, large leaflet size, large 2-seeded pods with
naturally occurring no constriction, pronounced beak, medium large flattened seeds, pink testa, 67% meat
pink variants in content, 45% oil content, field resistance to early leaf spot.
Makulu Red
Flamingo Virginia PI 2619111 Natal 1982

(bunch) Common
Plover Spanish Introduced from 1982

Brazil (PI 336954)
Swallo Virginia PI 2619111 Makulu 1982

(bunch) Pink selection
In the northern region, the main constraints to groundnut production are
early and late leaf spots, viruses (peanut stripe, peanut stunt, cucumber
mosaic, TSWV), aphids, Helicoverpa, Spodop:tera, thrips, nematodes and
drought. Surveys conducted in the 1970s indicated that more than
300000 ha were infested with nematodes in nine provinces of China.
Meloidogyne hapla is widespread in the north, whereas it is M. arenaria in
the south of the country. These nematodes cause on average 20-30% yield
loss in the country. Breeding began at the Peanut Research Institute at
Laixi in Shan dong Province in 1959. Since then 15 cultivars have been
released for cultivation in the province and other parts of the country
(Table 14.7). Following hybridization, the single-seed descent method has
been used to advance breeding generations. Twelve of the 15 cultivars
released by the institute are the result of hybridization and the remaining
three are pure line selections among local landraces. Hua 37 and Luhua 4
are very popular among farmers and have good export quality. Hua 37
covers more than 100000 ha in the country. With new production tech-
nology, which includes polyethylene mulching, these cultivars can produce
7.5 t pods/ha. The main emphasis in groundnut breeding in Shandong
Province has been to increase pod yield and improve quality. Quality
parameters that have received attention in breeding are large elongated
seed, high oil (55%), OIL ratio (> 1.4 for large-seeded virginia types, > 1.2
for spanish types), high protein (>30%), high blanchability, pink testa
colour, and flavour (by organoleptic test).
In the central region, early and late leaf spots, rust, bacterial wilt, viruses
(peanut stripe, peanut stunt, cucumber mosaic, TSWV) , aphids,
Helicoverpa, Spodoptera, thrips, leafhoppers, white grub, drought, water-
logging and high temperature are serious constraints to production. The
Oil Crops Research Institute at Wuhan is responsible for groundnut re-
search in Hubei Province in this region. The Institute maintains a collec-
tion of 4350 accessions of groundnut, including 130 wild Arachis species.
All accessions have been characterized for agronomic characters. In collab-
oration with the Peanut Research Institute in Shan dong Province, 4029
lines have been screened for resistance to nematodes (M. hapla) , to which
two lines - Tian Fu No.4 and Da Hua Cheng - have shown a high level of
resistance. Three other lines with moderate resistance and five lines with
tolerance also have been identified. Four thousand lines have been
screened for bacterial wilt, rust, late leaf spot and early leaf spot. Seventy
lines with resistance to bacterial wilt and many lines with resistance to rust
and late leaf spot have been identified, but a satisfactory level of resistance
to early leaf spot has not yet been located. Many of the lines resistant to
foliar disease were obtained from ICRISAT. The germplasm has also been
screened for biochemical factors. The protein content in the collection
ranges from 14.0% to 36.8% and oil content from 36.0% to 60.21 %. There
are many lines with OIL ratios greater than 3.0.
Breeding objectives at the institute include high yield, early maturity,
improved quality, and resistance to diseases and insect pests. Following
hybridization, the pedigree method is followed to advance breeding gener-
ations. From 1986 to 90, the significant achievements of the breeding group
at the institute included identification of sources of resistance to bacterial
wilt and rust.
About 200000 ha are infected with bacterial wilt in the central region of
China. Yield loss to this disease averages 10-15% and may go up to 60% .
Since 1970, more than 4000 germplasm accessions have been screened in
the field and screenhouse; 70 resistant lines have been identified. The
resistance in these lines is generally stable under field conditions but it can
break down under heavy artificial inoculation and with a highly virulent
strain. Inheritance studies involving spanish types indicated that resistance
to bacterial wilt is partially dominant and is governed by three major genes
with additive effects (Boshou et al., 1990).
Peanut stripe virus (PStV), although widely distributed in the country, is
mainly important in central and northern China. A 50% disease incidence
is often found in these areas, reaching up to 100% in many fields. In
southern China, the disease incidence is <1%. In laboratory and field
studies, 20% yield loss was observed with early infection of the virus. More
than 1300 germplasm lines have been screened without identifying any
resistant accessions.
In the central region mostly spanish and peruvian types are grown. Four
new groundnut cultivars have been released by the institute in the last five
years (Table 14.7). Current breeding activities (1991-95) include develop-
ment of cultivars with multiple resistance to diseases and pests, utilization
of wild Arachis species to develop cultivars resistant to leaf spot, screening
for resistance to virus diseases, screening for tolerance to acid soils and
breeding for increased nitrogen fixation.
In China's southern region, the primary constraints to production are
rust, bacterial wilt, waterlogging and soil acidity. Guangdong Province,
where mainly spanish types are grown, leads the region in groundnut
production and its Industrial Crops Research Institute, Guangzhou, carries
out groundnut research in the region. The main objective of its present
ground nut research programme is to develop high-yielding cultivars with
resistance to bacterial wilt and rust and adaptation to different growing
conditions in the province. The six sources of bacterial wilt resistance used
in the breeding programme are Teishan Sanliyue (a valencia cultivar from
China), Teishan Zhenzhu (a spanish cultivar from China), Xie Kong
Chung (a spanish cultivar from China), Schwartz (a spanish cultivar from
Indonesia), Yindu Huapi (a virginia cultivar from India) and Tianjin Don
(a runner cultivar from China). Sources of rust resistance have been
obtained from ICRISAT.
In Japan, groundnut is a minor crop. The main centre of production is
the Kanto region in the central part of the country. The consumption of
groundnut in Japan amounted to 85000 t in 1989, of which 44% was
TABLE 14.7 Groundnut cultivars released in East Asia
Cultivar Botanical type Pedigree Year of Characteristics

release
China (Northern Region)l
Fuhuasheng Spanish 1960
Hua 27 Virginia 1967
Hua 11 Spanish 1969
Hua 19 Virginia 1975
Hua 28 Intermed. 1979
between spanish
and virginia (M)
Hua 31 (Hai Hua 1) M 1984
Hua 37 M 1985 Good quality
Hua 39 (Luhua 4) Virginia 1986 Good quality
Hua 17 Virginia 1974 OIL ratio 1.45
Hua 98 Virginia 1974 Tolerant to drought
Luhua 3 Spanish 1982 High oil, resistant to bacterial wilt
Luhua 6 M 1986
Luhua 8 M 1988
Luhua 9 Virginia 1988 Good quality
China (Central Region)

El Hua 4 Hongmei Zhao I El High yield, early maturity, high quality, tolerant to drought
Hua2
Zhong Hua 1 El Hua 3 I Taishan High yield, tolerant to leaf spot
Zenghou
Zhong Hua 2 El Hua 4 I Taishan High oil and protein, resistant to bacterial wilt
Sanlirou
Zhong Hua 117 Resistant to rust, moderately resistant to bacterial wilt, high protein, high
yield
TABLE 14.7 Cant.

release
China Southern Region)

Yue You 39 Yue You 1161 Yindu Resistant to bacterial wilt and rust
Huapi
Yue You 223 Shan You 261 EC Tolerant to rust
76446 (292)
Yue You 92 Yue You 1161 Xie Resistant to bacterial wilt, high in oil content (54%)
Kang
Yue You 256 Yue You 1161 Xie Resistant to bacterial wilt, high yield
Japan
Wase-dairyu Spanish Early maturing, large seed
Tachi-masari Spanish Early maturing, large seed
Chiba-handachi Medium maturing cultivar with large seed
Nakate-yutaka Medium maturing, high yielding cultivar with good eating and external
quality
Azuma-yutaka Medium maturing, high yielding cultivar with good eating and external
quality
Sayaka Medium maturing (later than Nakate- Yutaka), high yielding, better suited
for roasting due to its thicker shell than Nakate-yutaka
Yude-rakka Early maturing, good eating quality, white pod colour with superior
external appearance, suitable for unshelled whole pod or frozen boiled
groundnut trade
TABLE 14.7 Cant.

release
Korea
Y ounghotangkong Virginia 1980 Late maturing, large elongated seed, pods with deep constriction
Saedltangkong Inter. between 1983 Early maturing, erect plant type, an intermediate type between spanish
spanish and and valencia
valencia
Jinpungtangkong 1986 Early maturing, small seeded, high yielding spanish type
(ICGS 35)
Daekwangtangkong Intermed. Florigiant I Chiba- 1985 Early maturing, high yielding, high oil content, erect plant type with few
between spanish handachi II Chiba- branches and large seed.
and handachi 131
valencia F393-6-3-2-3-1-2
Namdaettangkong Virginia (bunch) Virginia bunch 1988 Large seeded, high yielding, high in oil content, tolerant to Phoma
Improved I Suwoen 30 arachidicola
1 Cultivars released by the Peanut Research Institute, Laixi, Shandong Province, China.
produced locally and the rest was imported. Since the end of World War II,
groundnut breeding in Japan has pursued two main objectives: breeding
early-maturing cultivars for warm and cool areas, and breeding medium
and late-maturing cultivars. Because groundnut is a delicacy in Japan, both
eating quality and external quality are important attributes (Gocho, 1991)
and improvement in quality has received the most attention in groundnut
breeding. Sucrose content and hardness of seed are closely related with
eating quality and they decrease if harvesting is delayed. The seed hardness
is measured when the moisture content in seed is in the range of 5-9%.
(All cultivars under test should have the same moisture level within this
moisture range.)
Groundnut is also a minor crop in Korea, where yields are affected by
leaf spots, rust, and low temperature at the ripening stage. The main
breeding objective at the Crop Experiment Station, Rural Development
Administration, Suwan, is to develop cultivars with large seed and erect
plant type (Lee et ai., 1986, 1989).
14.4.3 Southern Asia

Groundnut research and production in southern Asia are dominated by
India. Other groundnut-growing countries in the region are Bangladesh,
Bhutan, Myanmar, Nepal, Pakistan and Sri Lanka. Except for Myanmar,
groundnut production in these countries is small. The crop in India and
Myanmar is grown mainly for edible oil production and in other countries
in the region for direct consumption or for use in confectionery. The region
accounts for 43.4% of the area and 35.7% of the production of groundnut
in the world. However, the average productivity in the region (0.94 t/ha)
remains below the world average (FAO, 1990). The main biotic constraints
to increased groundnut production in the region include diseases -late leaf
spot, rust, early leaf spot, collar rot (Aspergillus niger), stem rot (Sclero-
tium roifsii), A. flavus, bud necrosis disease (BNV) - and insects (thrips,
jassids, aphids, leaf miner, Spodoptera, Heiicoverpa, red hairy caterpillar,
whitegrub and termites). Abiotic constraints include drought, lack of high-
yielding cultivars adapted to local growing conditions, lack of availability
of good quality seeds and lack of small-scale farm machinery for groundnut
cultivation.
Introduction and reselection in introduced populations continue to be
the main methods of crop improvement in the region - with the exception
of India where, over the past decade, the majority of new cultivars have
resulted from hybridization between parents selected for their desirable
characteristics. However, in countries where the research programmes are
small and the scientists are responsible for more than one crop, depen-
dence on the introduction of improved germ plasm from various sources is
heavy. ICRISAT has played an important role in such introductions.
Prior to 1980, breeding efforts were directed mainly towards improving
TABLE 14.8 Groundnut cultivars released in South Asia

release
India (old popular cultivars)
Gangapuri Valencia Early maturing, 3-4-seeded, small-medium pods, preferred as table

variety, good source for earliness, popular in central India
Spanish Improved Spanish Selection from spanish 1905 Small-seeded, suitable for light soils (proposed for denotification)
peanut
Kopergaon I Virginia Selection from local 1933 Medium sized pods (proposed for denotification)
(bunch) variety
TMV2 Spanish Selection from 1940 Widely adapted, well suited for rainy and summer season cultivation in
'Gudhiatham Bunch', a southern India, a leading spanish variety in the past, still continues to be
local variety popular with farmers
AK 12-24 Spanish Selection from local 1940 Widely adapted, suited for medium to heavy soils (proposed for
variety de notification)
Punjab Groundnut I Virginia Selection from 1953 Wide adaptability (proposed for de notification)
(runner) 'Samrala Local'
Karad 4-11 Virginia Selection from local 1957 Late maturing, 1-3-seeded medium to long pods
(runner) variety
RSB 87 Virginia Selection from a 1961 3-seeded pods with dark red seeds
(bunch) Brazilian collection
J 11 (SB XI) Spanish Ah 4218 / Ah 4354 1964 Widely adapted, resistant to collar rot and A. flavlls seed colonization
S 206 Spanish Selection from 'Manvi 1969 Reticulated pods with slight beak and constriction
Local'
S 230 Virginia Selection from 'Tandur 1969
(runner) Local'
M13 Virginia Selection from NC 13 1972 Large-seeded variety with tolerance to leaf spots
(runner)
JL 24 Spanish Selection from EC 1978 Large dark green leaves, smooth 2-3-seeded pods, early in maturity
94943
TABLE 14.8 Cont.

release
India (cultivars released since 1980)
Kisan Spanish Spanish Improved B 31 1980 Small pod with prominent reticulation, released for Orissa State
M37 Virginia All C 6--4-7-2 1980 Two-seeded pods with small beak, light brown seed coat, released for
(runner) Punjab State
KRG1 Spanish Selection from 1981 Two-seeded medium sized pods, released for Karnataka State
'Argentine' variety
TG 17 Spanish 'Dark Green' Mutant I 1982 Large-seeded, pinkish seed coat, high harvest index, fresh seed dormancy,
TG 1 released for Maharashtra State
M 197 Virginia C 5011 U 4-7-2 1982 Dark green leaves, large-seeded pods with smooth reticulation, released
(bunch) for Punjab State
GG2 Spanish J111 1983 Two-seeded reticulated pods, early flowering with dark green leaves,
EC 16659 released for Gujarat State
Jawan Spanish 1111 Asiriya Mwitunde 1983 Medium elongated pods with moderate beak, rose seed coat, released for
Orissa State
CO2 Spanish EMS Mutant from 1983 Two-seeded medium plumpy pods with rose colour testa, released for
Pollachi 1 Tamil Nadu State
Dh8 Spanish Selection from RS 144 1984 Dark green leaves, compact plant, tolerant to late leaf spot, small pods
with smooth rose seeds round at one end and sharply pointed at the other,
released for Karnataka State
Chitra Virginia Spanish 5B-11 EC 1688 1984 Dark green leaves, variegated testa with rose background, released for
(runner) Uttar Pradesh State
Kaushal Virginia Selection from T 28 1984 2-3-1-seeded pods, compact plant with dark green leaves early in
(bunch) maturity, released for whole of India
UF 70-103 Virginia Introduction from USA 1984 Suitable for summer cultivation in Maharashtra State
(bunch)
GG 11 Virginia M 131 Gaug 10 1984 Leaflets and pod bigger than Gaug 10, released for Gujarat State
(runner)
TABLE 14.8 Cont.

release
TG3 Spanish A mutant of Spanish 1985 Spanish, medium-large pods, suitable for both rainy and summer seasons,
Improved tolerant to pod borer
MA 16 Virginia Selection from EC 1986 Large seeded suitable for HPS trade
(bunch) 16664
SG 84 Spanish Selection from ICGS 1 1986 Mainly 2-seeded, medium sized pods, suitable for summer/spring
cultivation in north India
M335 Virginia M13/F7 1986 2-1-3-seeded large pods with prominent reticulation and moderate
(runner) constriction, seeds large with light brown testa, large dark green leaves
with compact plant, released for Punjab State
ICGS 11 Spanish Selection from natural 1986 2-seeded smooth medium sized pods with no beak and slight to moderate
(ICGV 87123) hybrid population of constriction, seeds tan colour, 100-seed mass 60 g, oil 49%, protein 22%,
Robut 33-1 above average tolerance of end-of-season drought, photoperiod
insensitive. Field tolerance of bud necrosis disease, adapted to post-rainy
season cultivation in India, performs well in West Africa also
VRl1 Spanish TMV 7/ FSB 7-2 1986 Large pods with deep constriction and prominent beaks
ALR1 Spanish Pollachi 2/ PPG 4 1987 Small dark green leaves, red testa, resistant to rust and late leaf spot
Girnar 1 Valencia Xl4-4--B-19B / NC Ac 1988 Early maturing, resistant to late leaf spot, rust, collar rot, and seed
17090 colonization by A. flavus, 2-3-seeded with reticulated, constricted and
beaked pods
ICGV 87128 (ICGS 44) Spanish Selection from natural 1988 2-seeded smooth small to medium sized pods with no or little beak, seeds
hybrid population of tan in colour, 100-seed mass 60 g, oil 49% , protein 25%, field tolerance to
Robut 33-1 bud necrosis disease, good recovery from midseason drought, relatively
photoperiod inse'nsitive, adapted to post-rainy season cultivation in India,
performs well in Pakistan also
RG 141 Spanish Robut 33-1 INC Ac 1989 Spanish with dark green foliage suitable for black soils
2821
VRI2 Spanish JL 24 / CO 2 1989 Mostly 2-seeded large pods with moderate beak, constriction and
reticulation. Seeds light rose in colour, 100-seed mass 50 g, oil 48%
TABLE 14.8 Cant.

release
ICGV 87141 (ICGS 76) Virginia TMV 10 I Chico 1989 Mainly 2-seeded medium sized pods with moderate to prominent
(bunch) reticulation, slight to moderate constriction and beak, seeds tan in colour,
100-seed mass 44 g, oil 43%, protein 20%, good recovery for pod yield
from midseason drought, field tolerance to bud necrosis, adapted to rainy
season cultivation in India, performs well in Sudan also
ICGV 87121 (ICGS 5) Virginia Robut 33-1 I NC Ac 1989 2-seeded medium sized pods with none to slight beak and reticulation,
(bunch) 316 slight to moderate constriction, seeds tan in colour, seed mass 38 g/lOO, oil
58%, protein 22%, shows good recovery for pod yield from midseason
drought, adapted to rainy season cultivation in India
ICGS 1 (ICGV 87119) Spanish Selection from natural 1990 Mainly 2-seeded medium sized pods with slight to moderate constriction,
hybrid population of none to slight beak, and smooth to slight reticulation, seeds tan in colour,
Robut 33-1 oil 51.1 %, protein 21 %, 100-seed mass 35 g, shows good recovery for pod
yield from midseason drought, field tolerance to bud necrosis
Birsa Groundnut-3 Virginia Early Runner I Asiriya Early maturing
(bunch) Mwitunde
ICGV 87187 (ICGS 37) Spanish Selection from natural 1990 Mainly 2-seeded medium sized pods with slight reticulation, slight to
hybrid population of moderate constriction and none to slight beak, seeds tan in colour, 100-
Robut 33-1 seed 53 g, oil 48%, protein 23%, tolerance to end-of-season drought, field
tolerance of bud necrosis disease, photoperiod insensitive, tolerant to rust
and late leaf spot, adapted to summer season cultivation in India, also
performs well in Pakistan
ICGV 87160 Spanish Ah 63 I NC Ac 17090 1990 2-seeded stubby pods with moderate to prominent ridges, slight
[ICG(FDRS)lO] reticulation, beaks and constriction either absent or less conspicuous,
seeds tan in colour, 100-seed mass 36 g, oil 48%, protein 27%, resistant to
rust, tolerant to late leaf spot, field tolerance to bud necrosis disease, less
susceptible to stem and pod rots caused by S. rolfsii, moderately resistant
to leaf miner
VRI3 Spanish J 11 I Robut 33-1 1990 Small-seeded pods with moderate constriction and little or no beak
RSHY1 Spanish GDM/TMV2 1990 Suitable for residual moisture situation
ICGV 86590 Spanish X14-4-B-19-B I PI 1991 3-seeded pods, resistant to rust, tolerant to late leaf spot, bud necrosis
259747 disease, stem and pod rots, and Spodoptera
TABLE 14.8 Cant.

release
Pakistan
Banki Virginia (bunch) Introduction in 1973 160-180 days to maturity
No. 334 Virginia 180-200 days to maturity
(runner)
BARD 669 Spanish A composite of ICGS 1989 150-160 days to maturity, high yielding, high in shelling turnover
44 and ICGS 37
Bangladesh
Dhaka-l (Maizchar Spanish 1976 Oil 48-50%, shelling 75%, matures in 135-140 days, highly susceptible to
Badam) leaf spots
DG 2 (Basanti Badam) Virginia (bunch) - 1979 Mainly 2-seeded large pods, 170-175 days in maturity, seed dormancy for
40-50 days, tolerant to leaf spots
DM1 Valencia Introduced from India 1987 Very dwarf, early in maturity, tolerant to leaf spots and rust
Acc 12 Valencia 1988 Tolerant to drought, leaf spots and rust
Sri Lanka
Red Spanish Valencia 1961 Semi-erect, large 3-seeded pods, dark pink seed colour, 100-seed mass
45 g, shelling 68%, 110-120 days maturity
MIl Spanish 2-seeded medium pods with pink colour seed, 100-seed mass 40 g, shelling
72%, 110-120 days maturity
No. 45 Spanish Introduction from 1982 2-seeded medium pods with pink seed colour, 100-seed mass 45 g, shelling
ICRISAT, India 75%, 110-120 days maturity
Xl4-4-1-6-19-6 Spanish Introduction from India 1982 2-seeded medium pods with pink seed colour, 100-seed mass 48 g, 115-120
days maturity
TABLE 14.8 Cont.

release
Nepal
B4 Virginia (bunch) Introduction from 1976 135-140 days to maturity, 3-2 seeded medium sized pods, tan colour seed
Pakistan with high oil content, oil purpose cultivar
Janak Virginia NC Ac 343 1987 140--145 days to maturity, 2-seeded large pods, tan colour seed with high
(runner) oil content, moderately resistant to disease and insects, a dual purpose
cultivar
Myanmar
Sinpadetha 2 Spanish JL 24 1984/85
Sinpadetha 3 Virginia (bunch) Robut 33-1 1984/85
SP 121 Spanish 2-seeded small pods, early maturing
Magwe 10 Spanish SP 12110701 S 550-05 2-seeded small pods, high shelling (76%), high oil (54%), early maturing
Magwe 11 Spanish Selection from Shawat 2-seeded small-medium pods with high oil content (55%)
2116
Magwe 12 Spanish 2-1-3-seeded medium sized pods with high oil content (55%)
Magwe 15 Spanish UPL Pn-2 1 Kyaung 2-1-3-seeded medium pods, high shelling (77%), high oil content (54%),
Gone seed dormancy for 2 weeks
Kyaung Gone Virginia (bunch) - 2-1-seeded, seed dormancy up to 2 months
MS2 Virginia 2-3-1-seeded pods, seed dormancy up to 3 months
(runner)
Bhutan
In Bhutan some undefined cultivars are grown in small pockets in the
valleys for local consumption
yield potential. With the identification of sources resistant to major dis-
eases and insect pests at ICRISAT and in the national programme in India,
resistance breeding received a strong stimulus resulting in release of cuIti-
vars with multiple resistances in India. A genetic gain of 1.3-3.2% per
annum was achieved during the 1980s under rainfed conditions in India
(Nigam et at., 1991). A large number of cultivars have been released in
India, particularly since 1980 (Table 14.8). Notwithstanding the release of
several improved cuItivars, some very old ones are still grown extensively
due to lack of availability of seed: only 20% of the seed requirement in
improved cuItivars is met at present in India. The situation is not very
different in other countries of the region. Pakistan, Sri Lanka, Nepal, and
Bangladesh have very small groundnut research programmes and rely
mainly on introduction for improved germplasm. Although Myanmar has a
sizeable area under groundnut, its research programme is hampered by
lack of trained scientific manpower.
Approximately 80% of India's groundnuts are grown in the rainy season
(July-October). The remaining 20% is grown with irrigatiqn in the post-
rainy season (October/November-March/April) and the summer (January/
February-April/May). The groundnut area in the post-rainy/summer
season has increased recently as pod yields are high at this time. Varietal
requirements of rainy and post-rainy/summer seasons differ because of
differing disease and insect pest complexes occurring in them. High pod
yield, high shelling percentage and high oil content are requirements
common to both growing seasons. Additional requirements of improved
cuItivars in the rainy season are: drought tolerance; adaptation to agroeco-
logical zones differing in rainfall pattern and length of growing season;
fresh seed dormancy in spanish/valencia types; tolerance to insect pests
such as aphids, jassids, thrips, leaf miner, Spodoptera and white grub; and
tolerance to diseases such as early and late leaf spots, rust, collar rot, stem
rot, A. fiavus and bud necrosis. In the post-rainy/summer season, disease
pressure is generally very low but tolerance/resistance to insect pests such
as leaf miner and Spodoptera, tolerance of low temperature in the early
stages of crop growth, early maturity, and responsiveness to fertilizers and
irrigation are needed in new cuItivars.
Much of the emphasis in the past in groundnut breeding in India was
placed on the improvement of pod yield. The quality characteristics which
received attention included shelling percentage and oil content. Oil quality
itself received virtually no attention. During the VIn Plan (1990-95),
India's most recent programme for the improvement of agricultural pro-
duction, the following breeding activities have been accorded high priority:
• For dryland conditions, emphasis is on development of drought-
tolerant, high-yielding, early-maturing spreading groundnut cuItivars.
• For use in paddy fallows, early-maturing bunch cuItivars able to extract
residual soil moisture are being developed.
• For post-rainy/summer season irrigated conditions, the objective is to
produce high-yielding spanish cultivars tolerant of iron chlorosis.
• For rainfed crops, resistance to foliar diseases is a high priority.
There is also demand for cold-tolerant, early-maturing cultivars possessed
of fresh seed dormancy. High oil content is a primary objective for cultivars
developed for use as oilseeds, while large seeds and less susceptibility to
Aspergillus species are the objectives in cuItivars for confectionery.
For each breeding activity, targets have been fixed and responsibilities
have been assigned to main groundnut research centres under the aegis of
the All-India Coordinated Research Project on Oilseeds (AICORPO) at
the Indian Council of Agricultural Research (ICAR), New Delhi.
Hybridization between adapted cultivars and donor parents of desirable
characteristics, followed by selection for such traits combined with high
yield in segregating populations, has been adopted to achieve the target of
the breeding activities listed above. Wherever required, interspecific
hybridization is also being pursued. Some of the sources of desirable
characteristics in use in hybridization are:
For earliness:
Chico JB(E)559
TG(E)l ICGS 6
TG(E)2 ICGS 51
VG(E)55 ICGV 86309
91176 ICGV 86315
ICGS(E)21 ICG 11199
ICGS(E)22 CSMG 881
ICGS(E)52 CSMG 902
ICGS(E)217 CSMG 905
TG7 CSMG 917
J(E)5 CSMG 918
J(E)6 CSMG 9102
JB(E)194 Kadiri 3.
JB(E)262
For drought tolerance:
ICGV 86607 ICGV 87264
ICGV 86707 Gujarat Narrow Leaf Mutant, A l3
ICGV 87259
For cold tolerance:
A. monticola NRCG 9608
NGCG l339 CGC 498
For seed dormancy:
Dh8 TG7
CGC7 TG9
ICGS 30 TG 17
ALG56 C 390
Kadiri 3 CGC3
TMVlO RSHY6
For high shelling percentage:
J13 CSMG 916
Spancross Kadiri 3
For bold seed:
ALG62 CSMG 81-1
JSP(HPS)19 CSMG 83-1
Somnath CSMG 9101
CSMG 33 M13
CSMG 35
For high oil content:
NC Ac 17500 TMV 10
C 174 TG7
TMV3
For iron chlorosis tolerance:
NGS7 GG2
JL24
For resistance to rust and late and early leaf spots:
PI 259747 ICG(FDRS)69
PI 270934 ICGV 86350
PI 393516 ICGV 86598
PI 393517 ICGV 86707
PI 393643 ICGV 87160
PI 393527 ICGV 87261
PI 414331 ICGV 87264
NC Ac 17090 ICG 1697
NC Ac 17129 ICG 7894
ICG(FDRS)43 CSMG 84-1
ICG(FDRS)68
For tolerance to bud necrosis disease:
ICGV 86031
For insect tolerance:
Leaf miner:
ICG 5240 ICG 11786
GBFDS 273 ICGV 86137
GBFDS 592
Spodoptera:
ICGV 86350 ALG50
ICGV 87264
Jassids:
NC Ac 2663
Multiple insect resistance:
ICG 2271 JL 116
JL 83
For A. flavus tolerance:
Monir 240-30 J11
UF 71513 PI 337409
Ah 7223 PI337394F
Indian scientists have attempted to access genetic variability in the wild

relatives of groundnut. Interspecific hybridization between the tetraploid
A. hypogaea and diploid wild species A. cardenasii, A. stenosperma and
A. chacoense has been carried out in Tamil Nadu state in India.
Derivatives of the interspecific hybridizations are currently under evalu-
ation. Irradiation and chemical mutagens have been used frequently in
India to create additional variability for use in breeding programmes.
Cultivars such as MH 2, TG 1 (Trombay Groundnut 1), TG 3, BG 1 (Birsa
Groundnut 1) and BG 2 were developed by mutation breeding using
irradiation, and CO 2 (Coimbatore 2) from chemical mutagenesis.
14.4.4 South-east Asia

Groundnut is an important food legume and oil crop in south-east Asia.
Indonesia, Vietnam, Thailand and the Philippines are the major producers
in the region; other countries - Malaysia, Laos and Cambodia - have only
small areas under groundnut. Thailand, Vietnam and Indonesia are able to
meet their domestic demand but in the other countries there is a big gap
between domestic production and demand. Consumption of groundnut
pods and seeds in the boiled form is very popular in this region. Peanut
butter is a popular groundnut product in Malaysia and the Philippines.
The region grows groundnut on about 0.92 million ha with a total
production of 950000 1. Average pod yields are low compared with China
and the USA. Major production in the region comes from upland areas,
where groundnut is generally grown as a monocrop. In plantation areas it is
intercropped with young rubber, oil palm and coconut trees. A sizeable
area of groundnut is grown in rice fallows under residual moisture
conditions. .
Several biotic and abiotic factors are responsible for low productivity in
the region. The major constraints to increased groundnut production are
late leaf spot, rust, sclerotium wilt, bacterial wilt, peanut stripe virus, leaf
miner, leafhopper, Spodoptera, Helicoverpa, aphids, thrips, drought, acid
soils, low soil fertility, shade under plantation crops, low price of ground-
nut and lack of seed availability of improved cultivars.
Groundnut research in Thailand, Vietnam, Indonesia and the Philip-
pines is very active. Malaysia has a small groundnut research programme.
TABLE 14.9 Improved groundnut cultivars released in South-east Asia

release
Indonesia
Gajah Spanish Schwartz 21 f Spanish 1950 Adapted to upland cultivation, 95-100 days to maturity, seed size
45-50 gf100, pod yield 1.51 tfha, resistant to bacterial wilt
Kidang Spanish Schwartz 21 f Small 1950 Adapted to upland cultivation, 95-100 days to maturity, seed size
Japan 45-50 gf100, pod yield 1.5 tfha, resistant to bacterial wilt
Macan Spanish Schwartz 21 f Spanish 1950 Adapted to upland cultivation, 95-100 days to maturity, seed size
Banteng Spanish Schwartz 21 f Spanish 1950 Adapted to upland cultivation, 95-100 days to maturity, seed size
Pelanduk Spanish Kidang f VB 1 1983 Adapted to upland cultivation, 95-100 days to maturity, seed size
45-50 gf100, pod yield 1.5 tfha, resistant to bacterial wilt and A. fiavus
Tapir Spanish Kidang f VB 1 1983 Adapted to upland cultivation, 95-100 days to maturity, seed size
45-50 gil 00 , pod yield 2.0 tfha, resistant to bacterial wilt and A. fiavus
Tupai Spanish US 26 f Kidang 1983 Adapted to upland cultivation, 95-100 days to maturity, seed size
45-50 gf100, pod yield 2.0 tfha, resistant to bacterial wilt and A. fiavus
Rusa Spanish Gajah f AH 223 1983 Adapted to upland cultivation, 100-110 days to maturity, seed size
35--40 gf100, pod yield 1.5 tfha, resistant to bacterial wilt and rust
Anoa Spanish Gajah f AH 223 1983 Adapted to upland cultivation, 100-110 days to maturity, seed size
35--40 gf100, pod yield 1.5 tfha, resistant to bacterial wilt and rust
Kelinci Valencia Acc 12 1987 Adapted to upland and lowland cultivation, 100-110 days to maturity,
seed size 40--45 g(100, pod yield 2.0 tfha, tolerant to bacterial wilt and
resistant to leaf spot
Jepara Spanish 1989 Adapted to lowland cultivation, 90-100 days to maturity, seed size
35--40 gf100, pod yield 1.2 tfha, tolerant to bacterial wilt
Landak Spanish Schwartz 21 f Spanish 1989 Adapted to upland and lowland cultivation, 90-95 days to maturity, seed
size 45-50 gIlOO, pod yield 1.8 tfha, tolerant to bacterial wilt
TABLE 14.9 Cant.

release
Mahesa Spanish PI 350680 I Kidang 1991 Adapted to upland and lowland cultivation, 95-100 days to maturity, seed
size 46 g/100, pod yield 2.0 t/ha, resistant to bacterial wilt, tolerant to rust
Badak Valencia FESR 12 ILocal Depok 1991 Adapted to upland and lowland cultivation, 95-100 days to maturity, small
seed size, pod yield 2.0 t/ha, tolerant to bacterial wilt and leaf spot
Biawak Spanish 1991 Adapted to upland cultivation, 90-95 days to maturity, medium seed size,
pod yield 2.0 tlha, resistant to bacterial wilt
Komodo Spanish 1991 Adapted to upland cultivation, 95-100 days to maturity, medium seed size,
pod yield 2.0 t/ha, resistant to bacterial wilt and rust
Vietnam
Do Bac Giang Spanish Local cultivar
Sen Nghe An Spanish Local cultivar
Moket Spanish Local cultivar
Ly Spanish Local cultivar
Giay Spanish Local cultivar
Cuc Nghe An Spanish Local cultivar
Sutuyen Spanish Selection from China 1970
Tram Xuyen Spanish Selection from China 1970
V79 Spanish X-ray mutant of Bachsa 1980
77
B 5000 Spanish X-ray mutant of Bachsa 1983
77
Sen Lai (75-23) Spanish Sen Nghe An I Tram 1985
Xuyen
TABLE 14.9 Cant.

release
Philippines
UPLPn 6 CES 103 / PI 298115 1986
UPL Pn 8 CES 101 / PI 298115 1989
BPI Pn 2
UPLPn 2 Spanish Moket 1976
UPLPn4 Valencia Acc 12 (PI 314817) 1978
BPI P9 Spanish E.G. Red / Fante 17 1973
CES 101 Spanish Pure line selection from 1973
unknown cultivar
Thailand
Khon Kaen 60-1 Spanish Moket
Khon Kaen 60-2 Valencia TMV3 1988
Khon Kaen 60-3 Virginia Selection from NC 7 1988
Lampang Valencia
SK 38 Valencia Selection in local
cultivar
Tainan 9 Virginia (bunch) Introduction
Malaysia
MKTI 1990
Not much is known about Laos and Cambodia. The Peanut CRSP of
USAIO in Thailand and the Philippines, ACIAR of Australia in
Indonesia, and IORC of Canada in Thailand have supported or continue to
support groundnut research in the region. ICRISAT has played an import-
ant role in introducing improved germplasm in the region. In Thailand and
the Philippines, the national ground nut programmes have strong multidis-
ciplinary teams of scientists. In addition to introducing improved germ-
plasm, hybridization has been commonly adopted to develop new cultivars
in Indonesia, Thailand and the Philippines. Several cultivars have been
released in the region (Table 14.9). In Indonesia, almost all improved
cultivars are either resistant or tolerant to bacterial wilt; Schwartz 21, the
first disease-resistant groundnut cultivar developed through hybridization,
was released here as early as 1927.
Groundnut research activity in Malaysia is very limited. Improved germ-
plasm introduced from ICRISA T and other sources is evaluated for local
adaptation, including resistance to prevailing diseases and insect pests.
The research programme in Vietnam is in its infancy and suffers from
lack of trained manpower, poor infrastructure and paucity of resources.
However, in collaboration with ICRISA T, breeding activities covering
resistance to foliar diseases (late leaf spot and rust) and bacterial wilt,
earliness, high yield and improved seed quality have been initiated re-
cently. ICRISAT is developing single-seed descent breeding populations
derived from crosses between Vietnamese cultivars and other desirable
donor parents at its centre in India: at the Fs stage, these populations will
be grown in Vietnam for in situ selection.
In Indonesia. the main objective of the groundnut improvement pro-
gramme is to improve yield potential and adaptation to varying agroeco-
logy and cropping systems. The specific issues that receive attention are
early maturity, tolerance to excess soil moisture, tolerance to drought,
tolerance to soil acidity, tolerance to mineral toxicities, adaptation to inter-
and mixed-cropping, tolerance/resistance to insect pests and diseases, and
tolerance to heat. A massive field screening exercise was undertaken in
Indonesia to evaluate Arachis germplasm for resistance to peanut stripe
virus. No resistance was found among 9000 lines of A. hypogaea; among 54
accessions of wild Arachis species, only A. cardenasii was immune. A few
others showed a resistant reaction.
The primary objective of groundnut breeding in the Philippines is to
develop groundnut cultivars with desirable agronomic traits such as high
yield, early maturity, acceptable quality and resistance to rust, late leaf
spot, A. flavus, leafhopper and spider mites. In addition, the improved
cultivars should have tolerance/adaptation to drought, partial shade
and acidic soil conditions, and improved nitrogen-fixing ability. From
the screening activities, several promising sources of desirable characters
have been identified for use in the breeding programme (PCARRO,
1985).
They are:
Rust: Multiple insect pests:
PI 259653 NC Ac 343
PI 109839 NC Ac 2214
ICGS 55 ICG(FDRS)11
Sclerotium wilt: Bhairwa
IPB Pn 82-71-27
IPB Pn 82-68-16
Local factors:
Drought: Shade:
Acc 847 UPL Pn 2
EG Pn 12 IPB Pn 12-14
ICGS(E)123
ICGS(E)120
Acid soils: High nitrogenase activity:
IPB Pn 24-2 RLRS5
IPB Pn 24-3 RLRS7
IPB Pn 26-4 IPB Pn 49-12
BPI P9 57-422
UPL Pn 4
The most active groundnut breeding programme in the region is that
of Thailand, the objectives of which include: high yield and earliness;
adaptation to after-rice, unirrigated condition and before-rice growing
conditions; resistance to foliar diseases (rust and late leaf spot); resistance
to A. flavus, A. niger and Sclerotium rolfsii; and large-seeded confectio-
nery and boiling-type cultivars. Significant progress is being made in
achieving these objectives;. Two cultivars were released recently, and
several breeding lines with good promise have been identified and are
under evaluation.
14.4.5 Australasia
The Australasian region is not very important from the perspective of
global groundnut production. Production in the region is dominated by
Australia, which provides high quality groundnuts for world trade during
the off-season for producing nations in the northern hemisphere. Major
constraints to increased production in Australia include the foliar patho-
gens (early and late leaf spots and rust); soil-borne diseases (Cylindrocla-
dium black rot, Sclerotinia blight, and A. flavus); and drought. Other
countries in the region include Papua New Guinea, Solomon Islands,
Vanuatu, Tonga, New Zealand and Fiji, all of which produce only limited
amounts of groundnut for local consumption.
Australia has the most active research programme in the region. Prior to
the programme of varietal improvement started in 1977-78, the primary
source of cultivars in Australia was introduction. A high degree of mechan-
ization permits widespread use of cultivars with spreading or runner
growth habits. Large-seeded virginia-type cultivars such as Shulamith and
NC 7 are preferred here. The spanish cultivar 'McCubbin' was released by
the Australian national programme, the goals of which are yield improve-
ment, quality maintenance (particularly shelf life), and resistance to foliar
diseases.
Other countries in the region do not have breeding programmes but still
rely exclusively on introduction for new cultivars. Recently, Papua New
Guinea and Fiji obtained advanced breeding lines from ICRISAT for
evaluation and in situ selection. Red-seeded spanish is the preferred type
grown in the Solomon Islands and Papua New Guinea.
14.4.6 North America

The United States is the largest producer of groundnuts in North America
and conducts the bulk of the ground nut research in the region. Collection,
maintenance and evaluation of groundnut germ plasm have been high
priorities in the USA. Placement of a full-time groundnut curator for the
national germplasm collection at Griffin, Georgia, has helped to organize
efforts in this area. During the last decade, breeders have identified
considerable germ plasm that can be used to improve the groundnut
(Wynne and Halward, 1989b). At the same time, collection expeditions
have continued to add to the diversity available for improv~ment of the
groundnut (Simpson, 1983, 1990).
Utilization of the wild species of Arachis to improve the cultigen has
been investigated in the USA by research programmes in North Carolina,
Oklahoma and Texas. Much of the research has been concerned with the
crossing relationships among the various species and with cultivated
groundnuts. Pathways for the transfer of genetic material from the species
to cultivated groundnuts have been established (Simpson, 1991; Stalker
and Moss, 1987). The progress of research in this area has been reviewed
recently (Wynne and Halward, 1989b; Stalker and Moss, 1987).
Cultivar development programmes at state experiment stations in
Florida, Georgia, Oklahoma, North Carolina, Texas and Virginia and at a
private company (formerly Gold Kist; now Agratech) released numerous
cultivars (Table 14.10). Over the past 10 years, these have broadened the
genetic base of the groundnut crop in the USA and provided sources of
pest resistance (Knauft and Gorbet, 1989). Knauft and Gorbet assessed the
genetic diversity among cultivars released by 1988 and concluded that the
genetic base had been broadened considerably since 1976. This broadening
has continued through additional cultivar releases since this report (Isleib
and Wynne, 1992).
Cultivars released for their pest resistance include NC 6 (southern corn
rootworm), NC 8C and NC 10C (cylindrocladium black rot), Va 8IB
TABLE 14.10 Groundnut cultivars released in the United States
Cultivar Market type Pedigree Year of

release
Florigraze* Rhizoma Selection from PI 118457 (Arachis glabrata Benth. cv. 'Arb', collected by W.A. Archer near Campo 1978
Grande, Brazil, in 1936)
Arbrook Rhizoma PI 262817 (Arachis glabrata Benth. collected by W.C. Gregory (Col. No. 9569) near Trinidad; Itapua 1985
Department, Paraguay, in 1959)
Dixie Runner Runner Small White Spanish 3x-11 Dixie Giant 1943
Virginia Bunch 67* Runner Selection from 'Virginia Bunch' obtained in 1941 from East Georgia Peanut Co., Bulloch Co., GA 1945
Southeastern Runner Runner Selection from 'Southeastern Runner' 1947
56-15*
Early Runner Runner Small White Spanish 3x-2 1 Dixie Giant 1952
Florispan Runner Runner Basse 1 Spanish 18-38, GA 207-3 /I Early Runner 1953
Georgia 119-20 Runner Southeastern Runner 1 Dixie Giant, 210-4 /I Virginia Runner 1954
Florunner Runner F334A-3-14 (Florispan sib) 1 F230-118-B-8-1 (Early Runner sib) 1969
Altika Runner F393-7-1 (NC-FLA 14 sib) 131 GA 119-20, Southeastern Runner 1 Dixie Giant, 210-14/1 Virginia 1972
Runner
GK19 Runner F334-3-5-5-1 (Florispan derivative) 1 Jenkins Jumbo, F393--6 /I F334-9 (Florispan sib) 1973
Tifrun Runner Florida Small Spanish 1 Dixie Giant, F231-51 141 F385-1-7-2, Pearl (F228) /I F68-74 Sr1-2, McSpan 1977
(F13, Small White Spanish) 1 Virginia Jumbo Runner (F14), F249-42-3-1/31 Jenkins Jumbo, T1645
(selection from F416) 1 T1861, selection made in 1966 from local virginia stock in Georgia, thought to
have arisen from a virginia x spanish hybrid)
GK7 Runner F334-3-5-5-1 (Florispan derivative) 1 Jenkins Jumbo, F393-2 /I GK 19 1982
Sunbelt Runner Runner F392-12-B-28 (Florigiant sib) 1 V A Bunch 67, A4-4 /I Florunner 1982
Sunrunner Runner F439-16-10-1-1 (Florunner component) /I UF393-7-1 (NC-FLA 14 sib), UF334A-3-5-5-1 (Florispan 1982
derivative) 1 Jenkins Jumbo
Southern Runner Runner PI 203396 (resistant to Cercospora arachidicola and Phaeoisariopsis personata) 1 Florunner 1984
Langley Runner FlorunnerlPI 109839 1986
TABLE 14.10 Cont.

release
Okrun Runner Florunner I Spanhoma 1986

Tamrun 88 Runner Goldin I (Wilson County Peanut Co., Pleasanton, TX) I Florunner 1988
Georgia Runner Runner Krinkle-leaf (var. vulgaris) I PI 331334 (,Criollo', var. hypogaea from Bolivia) 1990
MARC I Runner Early Runner I Florispan, F439-17-2-1-1 (Florunner sib) II F459B-3-2-4-6-2-2-1 (Early Bunch 1990
component)
Improved Spanish 2B* Spanish Selection from local Spanish made C. 1918 at Florence, SC
Spanish 18-38* Spanish Selection from farmers' spanish stocks
Spanish No. 146 Spanish Spanish introduction (Coll. No. 146) obtained from India by Tom Huston Peanut Co.
GFA Spanish* Spanish Selection from 'Small Spanish' obtained from a grower in 1930 1941
Spantex* Spanish Selection from farmers' spanish stocks 1948
Dixie Spanish* Spanish Selection from Spanish introduction (Coll. No. 146) obtained from India by Tom Huston Peanut Co. 1950
Argentine* Spanish Selection from PI 121070 (var. vulgaris) 1951
Spanette* Spanish Selection from Spanish 18-38 1959
Starr Spanish Spantex I PI 161317 (var. vulgaris obtained in 1947 from SaIto, Uruguay) 1961
Spanhoma* Spanish Selection from 'Argentine' 1969
Comet* Spanish Selection from 'Starr' 1970
Spancross Spanish Argentine (PI 121070-1) I PI 405933 (Arachis monticola) 1970
Tifspan Spanish Argentine (PI 121070-1) I Spanette 1970
Tamnut 74 Spanish Starr II TPL 647-2-5, Spantex I Arachis monticola 1974
Goldin I Spanish Obtained from E. Goldin, Faculty of Agriculture, Hebrew University of Jerusalem, Rehovot, Israel 1976
by the Wilson Co. Peanut Co., Pleasanton, TX
Toalson Spanish PI 221057 (var. vulgaris) I Selection 26 (Spantex sib), TPL 673-A II Starr 1979
Pronto Spanish Chico I Comet 1980
TABLE 14.10 Cant.

release
Spanco Spanish Chico I Comet 1981
Tennessee Red* Valencia Selection from farmers' valencia stocks
New Mexico Valencia Valencia Selection from 'New Mexico Valencia' 1971
A*
McRan* Valencia Selection from African plant introduction 1973
New Mexico Valencia Valencia Selection from PI 355987, irradiated 'Colorado Manfredi' obtained from the research station at 1979
C* Manfredi, Argentina
Georgia Red Valencia UF439-16-10-3 (Florunner component) I New Mexico Valencia A 1986
NC4* Virginia Selection from 100 plant isolations made in 1929 from NC farmers' cultivars by P.H. Kime, NCSU 1944
agronomist, Selection #4 deemed typical virginia bunch
Holland Jumbo* Virginia Selection from farmers' virginia stocks 1945
Holland Virginia Virginia Selection from farmers' virginia stocks 1945
Runner*
Virginia Bunch 46-2* Virginia Selection from 'Virginia Bunch Large' 1952
Virginia Bunch G2* Virginia Selection from 'Virginia Bunch' obtained in 1941 from East Georgia Peanut Co., Bulloch Co., GA 1952
Virginia Bunch G26* Virginia Selection from 'Virginia Bunch' obtained in 1941 from W.A. Groover, Bulloch Co., GA 1952
NC 1 Virginia NC4 I Improved Spanish 2B 1952
NC2 Virginia Basse I Spanish 18-38, GA 207~2 II White's Runner 1952
Virginia 56R * Virginia Selection from 'Atkins Runner' 1956
NC4X Virginia Selection from irradiated 'NC 4' 1959
Florigiant Virginia Basse I Spanish 18-38, GA 207-3 II F230-118-2-2 (same as F230), F334A-5-5-1 131 F359-1-3-14, 1961
Jenkins Jumbo II F230-118-5-1, Dixie Giant I Small White Spanish 3x-2
Virginia 61R* Virginia Selection from 'Atkins Runner' 1962
NC5 Virginia NC 1 II C12, PI 121067 I NC Bunch 1964
Shulamith Virginia Florigiant I F334A-B-17-1 (Florispan derivative) 1968
TABLE 14.10 Cont.

release
NC17 Virginia F334A-3-5-5-1 (Florispan derivative) I Jenkins Jumbo 1969

Virginia 72R Virginia VA 61R I VA A89-15 (selection from farmers' stocks, perhaps Atkins Runner) 1971
NC-FLA 14 Virginia Jenkins Jumbo I F334A-3-5-5-1 (Florispan derivative) 1973
Keel 29* Virginia Selection from 'Florigiant' 1974
Avoca 11* Virginia Selection from 'NC 2' 1976
GK3 Virginia Florida Small Spanish I Dixie Giant, F231-51 141 F385-1-7-2, Pearl (F228) II F68-74 Srl-2, McSpan 1976
(F13, Small White Spanish) I Virginia Jumbo Runner (FI4), F249-42-3-1 131 Jenkins Jumbo, F416-2
151 F392 (Florigiant sib)
NC6 Virginia NC Bunch I PI 121067, C12 II C37 (same as C12), GP-NC 343 (selection from NC Ac 4508) II VA 1976
61R: Resistant to SCR
Early Bunch Virginia Virginia Station Jumbo 141 F385-1-7-4, Pearl (F228) II F68-74 Srl-2, McSpan (F13, Small White 1977
Spanish) I Virginia Jumbo Runner (F14), F249-42-3-1 131 Jenkins Jumbo, F406A 151 F420, F231-51
(Dixie Runner sib) I F392-12-1-7 (Flori giant sib)
NC7 Virginia NC 5 II F393, F334-3-5-5-1 (Florispan derivative) I Jenkins Jumbo 1978
VA81B Virginia F392-8 (Florigiant sib) 131 GA 119-20, Southeastern Runner I Dixie Giant, 210-14 II Virginia Runner 1981
NC8C Virginia NC 2 II A48, NC 4 I Spanish 2B, NC Ac 3139 131 Florigiant 1982
NC9 Virginia NC 2 I Florigiant 1985
NClOC Virginia NC 8C I Florigiant 1988
NC-V 11 Virginia Florigiant INC 5 II Florigiant I PI 337396 (var. [astigiata) 1989
VA-C 92R Virginia Florigiant II F393, F334-3-5-5-1 (Florispan derivative) I Jenkins Jumbo, NC Ac 17213 131 NC 7 1992
VC-1 Virginia F334-3-5-5-1 (Florispan derivative) I Jenkins Jumbo, F393 II F334 (Florispan derivative) I F393 131 1991
F392 (Florigiant sib) I GA 186-28 II F439 (Florunner component)
* Developed by selection within a plant introduction or existing cultivar.

(sclerotinia blight) and Southern Runner (late leaf spot). The cultivars that
have been released primarily for their pest resistance have generally com-
promised one or more agronomic traits, making them less competitive in
absence of the pest.
Considerable effort to develop pest-resistant groundnut cultivars began
during the 1980s in the USA. Wynne et al. (1991) summarized progress in
breeding for disease resistance. They concluded that although several USA
breeding programmes had been initiated for resistance to diseases -
Aspergillus spp. (aflatoxin), tomato-spotted wilt virus, nematodes, early
and late leaf spots, sclerotinia blight, and cylindrocladium black rot - few
cultivars had yet been released, due to the short duration of the efforts.
However, many sources of disease resistance were identified by screening
programmes during the 1980s and breeding for disease resistance is now a
priority in most USA programmes. Much progress can be expected.
Considerable effort in the USA has also been devoted to the use of wild
species of Arachis for sources of resistance to pests. Programmes to
transfer the high levels of resistance or immunity to early and late leaf
spots, rust, nematodes and viruses were active during the 1980s (Stalker
and Moss, 1987; Wynne and Halward, 1989a). To date, no cultivar incor-
porating germplasm from diploid wild species has been released.
Recently the groundnut industry identified quality and aflatoxin resist-
ance as two major issues that needed additional research and were con-
sidered of highest priority because of the effect they have on the export of
groundnuts. Substantial funding from the National Peanut Foundation and
USDA has increased conventional breeding and molecular genetic
research to address these problems.
Several researchers in the USA are now investigating and developing
methodologies to use molecular techniques for groundnut improvement.
The use of RFLPs as molecular markers is being investigated by a
University of Georgia researcher (Kochert and Branch, 1990) in co-
operation with several others. Little variation has been reported among
cultivars but abundant polymorphism has been found among the diploid
species of Arachis. Similar results were found for isozymes (Grieshammer
and Wynne, 1990; Stalker et ai., 1990).
Several USA researchers are investigating somatic embryogenesis and
plant regeneration in groundnuts. At least two laboratories have devel-
oped a repetitive somatic embryogenesis system and have established
plants in soil (Durham et ai., 1991; A. Weissinger, North Carolina State
University, personal communication, 1991). These successes should ex-
pedite the use of gene transfer systems in the crop. The use of microprojec-
tile bombardment as part of a gene transfer system in groundnut is being
evaluated in at least two laboratories. The success of these systems will
allow the movement of agronomically important genes into the groundnut.
Several laboratories are identifying and sequencing genes from viruses
and from other plants that may be useful in improving the groundnut. This
research is receiving funding support from the Peanut CRSP, private
companies, the USDA and state experiment stations.
14.4.7 South America

The area under commercial groundnut production in South America is
about 350000 ha. Argentina ranks first in groundnut area in the region
(180000ha), followed by Brazil (100000ha) and Paraguay (30000ha).
The area in other countries, such as Bolivia, Chile, Ecuador, Peru,
Uruguay and Venezuela, does not exceed 5000 ha. Although the region'S
area under groundnuts has been declining, the total production has not
suffered significantly, due to increase in productivity: average yields in the
1980s were nearly 50% higher than those of the 1970s. New crop pro-
duction technology and improved cultivars have contributed to increased
yields.
Average seed yield in Argentina has increased from 0.79 t/ha in the
1970s to 1.20 t/ha in the 1980s. During the 1970s two valencia cultivars,
Colorado Irradiado INTA and Blanco Rio Segundo, contributed 80% to
the total groundnut production; the remaining 20% was contributed by
Blanco Manfredi 68, a derivative of a cross between virginia and spanish
types (Godoy and Giandana, 1992). Since then, the varietal picture in the
country has changed completely as virginia runner types proved better
adapted to the main groundnut-growing region of the country. By 1989,
Florman INT A and Florunner accounted for 80% of the total groundnut
area and production. This development is somewhat disturbing in view of
the international community's expressed desire to maintain genetic diver-
sity in food crops, particularly in the centres of diversity for those species.
However, it is necessary to balance against this desire the needs of the
individuals and nations in those regions. The good adaptation of runner
types to the climate and soil in the country led Argentina to become the
third largest exporter of edible groundnut in the world, after the USA and
the People's Republic of China.
Average pod yields in Brazil in rainy seasons are 2.0-2.1 t/ha and in the
dry season about 1.5 t/ha. The reduction in area and production has been
dramatic: the cultivated area in 1972 was 759000 ha and it declined to
100000 ha in 1988, while production fell from 956000 to 167000 t in the
same period. The main reason for such a sharp decline was establishment
of soybean as the leading oilseed crop in the country. However, the
average yield of groundnut has increased from 1.5 t/ha in the 1970s to
1.8 t/ha in the 1980s in Sao Paulo province (the main groundnut-growing
area in the country). In the Ribeirao Preto region, the pod yield averages
2.5 t/ha but yields up to 4.0 t/ha can be obtained with the red valencia
cultivar Tatu, which has a short growing cycle of 90-100 days and now
occupies 80% of the ground nut-growing area in Sao Paulo. Another
cultivar, Tatu Branco, which is similar to Tatu except for its seed colour,
occupies 10% of the groundnut area; it has undefined tolerance to drought
and is adapted to low fertility. Recently three new cultivars with 15-20%
higher yield than Tatu have been released - Tupa, Oira and Poitara. All
three are derived from valencia-by-spanish crosses; they mature in 110-120
days and have two-seeded medium sized pods.
Groundnut cultivation in Bolivia is manual and local cultivars are grown.
The three main local cultivars are Coloradito Palmer, an erect type with
125 days maturity; Cuero Padilla, a semi-erect type with 135 days maturity;
and Bayo Gigante (also called Colorado Grande), a runner type with 145
days maturity.
In Paraguay, groundnuts are grown in three regions which differ in soil
type, climatic conditions and level of technology input. In the Chaco
region, cultivation is mechanized and spanish cultivars are grown. In the
central region, valencia and spanish types are grown by small farmers in
less fertile soils with low levels of technology input. In the southern region,
long-season virginia types are cultivated on fertile soils. The present yields
in Paraguay (1.3 t/ha) are 50% higher than those of 20 years ago.
Pod yields in Uruguay range from 0.7-1.8 t/ha. Groundnuts are
generally cultivated by small farmers on acidic sandy soils which are low in
Ca content, with family labour and little technology. Predominantly valen-
cia types are grown; spanish and virginia types are also cultivated to a
limited extent.
In 1990, the southern nations of South America (Argentina, Brazil,
Bolivia, Paraguay and Uruguay) initiated a co-operative research effort
called PROMANI (pro = program; manf = groundnut). Its objective is to
promote groundnut research and extension activities in the participating
countries. Research in Argentina and Brazil has been intensified since the
early 1980s. Both countries have their own active breeding programme,
whereas other PROMANI countries rely mostly on introduction of
improved cultivars and'selection in local cultivars/landraces.
In Argentina, groundnut research is focused on studies of the taxonomy
of the genus Arachis and on the development of new cultivars of medium
duration (125-130 days) with tolerance to drought and leaf spots, resist-
ance to A. flavus infection, high OIL ratio, low iodine value, improved
content and quality of seed proteins, and improved flavour and aroma. In
Brazil, research objectives include germplasm collection and taxonomic
studies on genus Arachis; breeding for resistance to late leaf spots; selec-
tion of early-maturing, high-yielding, red-seeded valencia/spanish types
with improved pod/seed appearance and shelling out-turn; development of
high-yielding virginia runner cultivars with resistance to leaf spot and other
diseases; resistance to Aspergillus infection; and resistance to thrips.
Development of high-yielding, leaf spot-resistant cultivars with accept-
able agronomic and quality attributes will benefit the South American
region most. Other diseases which could be potentially important in the
region are rust, scab and Sclerotium. Except for some areas in Argentina,
Accomplishments and future efforts 615
groundnuts in the region are generally grown under rainfed conditions.
Drought is the most common abiotic stress in the region.
14.5 ACCOMPLISHMENTS AND FUTURE EFFORTS
On a worldwide basis, the most important results of groundnut breeding in

the past 10-20 years have been the identification of sources of resistance to
the three globally important foliar fungal pathogens and the transfer of
resistance into breeding populations with the locally appropriate agrono-
mic attributes. It remains to be seen whether release of cultivars resistant
to rust and leaf spot will significantly affect patterns of groundnut
production.
Closely following the foliar diseases in importance is the aflatoxin prob-
lem. Despite the identification of seed-coat resistance and the release of
IVSCAF-resistant cultivars, aflatoxin contamination remains the largest
single problem affecting international trade in groundnut. The recent
adoption by the European Community of extremely low tolerances for
aflatoxin may eliminate some nations from the array of groundnut
exporters. Although this problem is certainly not confined to the realm of
plant breeding, the international community looks primarily to a genetic
solution.
Breeding for resistance to insect pests has not been emphasized to the
same degree as breeding for resistance to foliar diseases. Common foliar
diseases occur with great regularity in most parts of the world while many
insect species require particular environmental conditions in order to reach
the population densities necessary to cause economic damage. Under
management systems with minimal or no application of insecticides and
fungicides, insect pest populations may be curbed by the presence of
predatory insects and animal or fungal parasites. Host, pests, predators
and parasites exist in a balance sensitive to subtle changes in the ecological
dynamic. In such production systems, pest resistance of low or intermedi-
ate level may be sufficient to shift the balance in favour of the host plant. In
many developing nations, the microeconomics of the production system
and the infrastructure for distribution and acquisition of pesticides prohibit
widespread use of pesticides for control of insects. Host-plant resistance to
insects will be the most effective means of reducing losses in yield and
quality associated with insect depredation.
Under intensive management systems, insect pests are controlled by
applications of pesticides that may also destroy beneficial species whose
absence allows unchecked growth of pest species that develop later in the
growing season, thereby necessitating further applications of pesticides.
Recently, socioeconomic forces in developed nations have created the
concept of LISA - low-input sustainable agriculture - as a paradigm for
mechanized agricultural production systems. These forces include demand
by consumers for agricultural products free from pesticide residues, public
concern over the effects of pesticides on the environment, reduction of
production costs, and the increasing difficulty encountered by manufac-
turers of pesticides in obtaining government approval for their registration
and sale. Key concepts of LISA include minimal application of pesticides
that have potentially harmful effects on consumers or environment; em-
phasis on soil conservation, including reduced tillage and use of green
manure animal waste as sources of organic matter and incorporation of
leguminous species into rotations to reduce use of mineral fertilizers that
can contaminate groundwater supplies. In short, LISA comprises a set of
production practices which by choice avoid extensive reliance on the
products of the chemical revolution that has so dramatically changed the
face of agriculture in developed nations in the last 50-60 years. While use
of herbicides and fungicides are affected by these practices, insecticides are
probably affected most because of their generally greater acute toxicity to
mammals. This trend may provide impetus for increased efforts in breed-
ing for insect resistance in developed nations. It remains to be seen
whether the consuming public in developed nations is sufficiently desirous
of pesticide-free produce to accept groundnuts bearing evidence of insect
feeding. Assuming that it is not, countries supplying edible groundnuts will
need to deploy cultivars with high levels of resistance to insects, a practice
that will certainly place strong selective pressure on pest populations.
Until recently, the gene pool for cultivated groundnut comprised the
global collection of the cultigen (some 12000 accessions) and the smaller
collection of Arachis species of which genes only from species of section
Arachis were accessible through sexual transfer. In the summer of 1992,
researchers from several public and private institutions reported success in
transforming groundnut with exogenous DNA and regeneration of fertile
plants from transformed tissues. Transformation has been effected through
microprojectile bombardment of embryonic axes (Brar et al., 1992),
embryogenic immature cotyledonary tissue (Weissinger et al., 1992) and
callus derived from embryonic leaflets (Weissinger et al., 1992), and
through electroporation of protoplasts (Demski et al., 1992). It would
appear that the array of transformation techniques effective in soybean can
be adapted to groundnut through modification of protocols. This effec-
tively converts the gene pool from a portion of the genetic information in
genus Arachis to virtually all genes in the planetary biosphere. The key
problem in groundnut breeding is changing from location of sources of
useful genes within the cultigen to identification of the genes per se, i.e. the
DNA base sequences, of potential economic value in groundnut regardless
of the source of those genes. Issues of the proprietary nature of such genes
and the payment of royalties, particularly by groundnut producers in
developing nations, will doubtless interest the ground nut breeding commu-
nity for decades to come.
References 617
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Proceedings of a Discussion Group Meeting, 24-28 September 1984, ICRISAT,
Sanders, TH. and Mixon, A.C. (1978) Effect of peanut tannins on percent seed colonization
and in vitro growth by Aspergillus parasiticus. Mycopathologia, 66,169-173.
Simpson, C.E. (1983) Plant exploration: planning, organization and implementation with
special emphasis on Arachis, in Conservation of Crop Germplasm: an International
Perspective, (eds W.L. Brown, TT Chang, M.M. Goodman and Q. Jones), Crop Science
Society of America Special Publication, No.8, CSSA, Madison, Wis., pp. 1-20.
Simpson, C.E. (1990) Collecting wild Arachis in South America past and future, ill IBPGR
International Crop Network Series. 2. Report of a Workshop on the Genetic Resources of
Wild Arachis Species Including Preliminary Descriptors for Arachis. IBPGRlICRISAT,
Rome, pp. 10-17.
Simpson, C.E. (1991) Pathways for introgression of pest resistance into Arachis hypogaea L.
Singh, A.K., Subrahmanyam, P. and Moss, J.P. (1984) The dominant nature of resistance to
Puccinia arachidis in certain wild Arachis species. Oleagineux, 39, 535-538.
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Smith, O.D., Boswell, T.E., Gricher, W.J. and Simpson, C.E. (1989) Reaction of select
peanut (Arachis hypogaea L.) lines to southern stem rot and Pythium pod rot under varied
disease pressure. Peanut Science, 16, 9-13.
Sowell, G., Smith, D.H. and Hammons, R.O. (1976) Resistance of peanut plant introduc-
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Stalker, H.T and Moss, J.P. (1987) Speciation, cytogenetics, and utilization of Arachis
species. Advances in Agronomy, 41, 1-40.
Stalker, H.T., Jones, T.M. and Murphy, J.P. (1990) Isozyme variability among Arachis
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Subrahmanyam, P., Ghanekar, A.M., NoIt, B.L. et al. (1985) Resistance to ground nut
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36,377-385.
CHAPTER 15
Utilization of Arachis species

as forage
B.C. Cook and I.C. Crosthwaite
15.1 INTRODUCTION
Legumes are no less important in the nutrition of grazing animals than

they are in human nutrition. They serve a dual role: firstly as a source
of high protein fodder in the animal diet, and secondly to inject nitrogen
into the forage system by virtue of the symbiotic relationship between
many leguminous species and the bacterial genera, Rhizobium and
Bradyrhizobium. Nitrogen input from the legume is one of the prime
determinants of the productivity of a mixed grass/forb pasture, and high
levels can only be achieved through maintaining a healthy, productive
legume component in the pasture. While nitrogen can also be supplied as a
constituent of chemical fertilizers, there is often farmer resistance to such
practice based on economic and environmental considerations. Legumes
vary in their capacity to fix nitrogen, as well as in their general suitability as
forages in terms of productivity, palatability, toxicity, growth habit and
adaptation to environment. Further, the legume flora varies considerably
from country to country and region to region. Where there is a natural
dearth of suitable forage legumes, researchers draw germplasm from
similar environments elsewhere in an endeavour to provide more pro-
ductive, stable pasture systems. Thus, traditional temperate legume genera
such as Trifolium, Medicago and Lotus have proved to be of less value in
the tropics and subtropics than they are in temperate and mediterranean
regions, necess,itating a search for, and evaluation of, a range of tropical
forage species.
The science of tropical forages is in its infancy in comparison with that of
temperate forages. It spans a period of little more than 50 years, during
which commercial forage legumes have been drawn from many genera, the
most widely planted probably coming from Centrosema, Macroptilium,
Stylosanthes and Desmodium. However, the genetic potential of these and
Arachis hypogaea 625
other genera, including Arachis, has been only partly exploited and evalu-
ation continues. The considerable contribution of Arachis to the pasture
lands of South America is well recognized (Hartley, 1949; Higgins, 1951b),
leading pasture plant evaluators to assess Arachis for pasture improvement
in Australia (Miles, 1949), India (Aiyadurai, 1959) and the USA (Prine,
1964; Blickensderfer et al., 1964; Beaty et al., 1968), resulting in the release
of a number of cultivars.
15.2 ARACHIS HYPOGAEA
While the commercial groundnut or peanut (Arachis hypogaea) has gained

greatest prominence by virtue of the quality and quantity of its seed, the
value of the vegetative portion of the plant, both fresh and as hay, has long
been recognized (Burke, 1850; Easby, 1851). The crop is grown in many
parts of the world, and in various ways makes a significant contribution to
animal production.
15.2.1 'Hogging off'

Whole groundnut plants have been grazed and rooted up by pigs for
generations. 'Hogging off' or turning pigs into the fields to root out the
nuts has been very popular in the United States. During the lO-year period
1925-34, approximately 35% of the total groundnut crop of 785000 ha was
hogged off rather than being harvested (Morrison, 1938). As late as 1973,
3% of groundnuts were used in the field for fattening pigs (Bogdan, 1977).
Groundnut crops which are hogged off have a beneficial effect on the soil
compared with those harvested conventionally (York and Colwell, 1951).
Corn and cotton yields increased after hogged off groundnuts, probably
due to the nitrogen returned to the soil.
15.2.2 Use of haulms

Haulms can be defined as that part of the plant remaining after the pods
have been removed. This includes leaves, stems and pegs. The amount of
root material remaining depends on the harvesting method.
(a) Asia
In northern China, up to 80% of the haulm may be stored for use as a
fodder for horses, pigs and sheep during winter, whereas only 50% is
stored in southern China where it is warmer and there is a lower need for
stored hay (Liao Boshu, personal communication).
South-east Asian farmers make significant use of groundnut haulms,
which are sold in bundles and cartloads during harvest time and are often
626 Utilization of Arachis species as forage
fed green. In Indonesia, horses carry fresh haulms for consumption while
away from home. In the Philippines, groundnut haulms are transported to
animal sheds away from the riverine production areas, which are subject to
inundation during the wet season (R.B. Santos, personal communication).
Dry haulms are fed to pigs and cattle in Vietnam, where the crop is
windrowed. However, few Thai farmers use groundnut haulms to feed
stock (V. Benjasil, personal communication).
(b) Africa
Groundnut haulms are an important feed source in many parts of Africa.
Haulms are commonly used in the northern part of Cameroon as cattle
feed and are actively traded during harvest (Essomba et al., 1990). They
are also used in Gambia (Drammeh, 1990), Nigeria (Misari et al., 1990)
and Ethiopia (A. Wakjira, personal communication), but feeding of
haulms to cattle is not yet widespread in the Republic of Guinea (Toun-
kara, 1990). While all of the haulm crop is fed to livestock in South Africa
(C.l. Swanevelder, personal communication), they are not used as stock
feed to any great extent by smallholder farmers in other southern African
countries because of spoilage by foliar disease and since the majority of
these smallholder farmers do not own livestock (G. Hildebrand, personal
communication) .
(c) India
Groundnut haulms are widely used in India as cattle feed (Nagaraj, 1988).
They are fed green or stored in 'hay stacks' and are often mixed with other
fodder materials. Farmers are happy with good haulm yields, even when
pod yields are poor, because their economy depends on milk animals.
(d) Australia
The sale of groundnut hay in southern Queensland can add significantly to
the income generated from the crop. Typically the value of the total crop
could be increased by 10-20% by selling the haulms. In some drought
years, crops have been baled and sold without threshing.
(e) USA
Widespread use was made of groundnut haulms in the United States.
Bogdan (1977) estimated that haulms from 20% of the peanut area or
100000 ha was utilized as hay. The value of goober pea (groundnut) vines
as stockfeed was recognized in the USA as early as 1851 (Higgins, 1951).
15.2.3 Haulm quality
Groundnut haulms are valued for their nutrient content, particularly their
high digestibility and protein. However, as the haulm component is only
secondary to the production of kernels, its quality varies considerably
(Table 15.1). In most situations the optimum time for haulm production
and quality would be well before the crop is harvested for maximum kernel
yields. Halevy and Hartzook (1988) found that the decline in nitrogen and
phosphorus levels in the stems and leaves began 64 days before harvest and
potassium levels 37 days later. The number of pods remaining in the hay
also influences quality (Table 15.2), particularly in relation to digestible
protein and metabolizable energy.
Weather conditions may affect haulm quality. A prolonged drought or a
long period of wet weather prior to harvest leads to leaf and stem deterio-
ration. The time that is most critical to haulm quality is between lifting and
removal of the haulms from the field: wet weather when the plants are
drying on the soil surface causes a rapid fall in quality.
Where mechanized threshing is used, the weather and bush condition at
threshing are important to haulm quality. If the humidity is very low, much
of the valuable leaf material is broken up into fine particles and lost when
passing through the thresher, leading to very stalky hay. Groundnut leaf
can contain more than twice the nitrogen or crude protein content of either
stem or root fractions (Powell, 1986). Some threshers in Australia have
been modified to leave the haulms in windrows for easy pick-up by
haybalers instead of spreading the discarded haulms evenly over the
paddock.
Disease is also a vital factor in haulm quality and quantity. Foliar
diseases such as leaf spots (Cercospora arachidicola and Phaeoisariopsis
personata) , rust (Puccinia arachidis) and some virus diseases can reduce
the leaf component significantly.
Quality can be influenced by plant population, through its influence on
leaf to stem ratios. Increases in plant popUlation significantly increase the
percentage of total dry matter partitioned to stems and decrease the
percentage partitioned to pods, while the partitioning to leaves does not
change (Kvien and Bergmark, 1987). Similar effects have been recorded
with planting dates, where a greater proportion of vegetative dry matter
was fixed in stems rather than leaves of early planted crops compared with
later plantings (Bell, 1986).
Significant differences among cultivars in leaf concentrations of potass-
ium, calcium and magnesium (Hallock and Martens, 1974) are of little
consequence in terms of feed quality.
15.2.4 Improving nutritive value

The quality of groundnut haulms can be changed by treating with an alkali
or urea. In vitro dry matter (DM) and organic matter digestibility were
TABLE 15.1 Nutritional value of post-harvest groundnut haulms
Source
1 2 3 4 5 6 7 8 9
Dry Matter (% ) 90 89-91 90-93 85 87-89 92.2

Crude protein (%) 6-20 13-14 19.9 8.3-15 12 9.6 9.6-12.1 14.6-15 9.3
Digest. protein (%) 2-6 7.3-13.2 5.7 6.5 4.9-6.1 7.6-7.9
TDN(%) 50 53 56 46.8-56.4
ME (MJ/kg) 8-11 9.8
Fat (%) 2 4.8 1.4-2.9 4.6 1.1
Ether extract (%) 0.8-2 2.3
NFE(%) 40.7 48.4 38.9-45.9 33.8
Carbohydrate (%) 42-46 38-47
Carbon hydrolysis (% ) 30.9
A-D-F (%) 38.4 40.3-43.1
N-D-F (%) 42
Crude fibre (%) 22-24 34.5 22-35 30.0 19 28-30.5 37.6
Ash (%) 5.7-6 9.9 11.9 9.1-15.3 9.1
Minerals (%) 9-17
Phosphorus (%) 0.7 0.42 0.3
Calcium 1.7 2.4 1.5
Sources: 1. W.J. Edwards, pers. comm. 2. Anon. (1982) 3. V. Benjasil, pers. comm. 4. Nagaraj (1988) 5. Shukla et al. (1985) 6. Ikhatua and
Adu (1984) 7. Combe lias et al. (1972) 8. Velasquez and Gonzalez (1972) 9. Leche et al. (1982)
TABLE 15.2 Nutrient value of groundnut hay of differing quality (after Bredon
et aI., 1987)
Good Average With some With pods

pods left
Good dry matter (% ) 90 92 91 92

Crude protein (%) 12.3 9.2 10.8 13.4
Digestible protein (%) 7.9 6.0 6.5 10.2
TDN(%) 50.0 49.0 62.6 71.6
ME (MJ/kg) 7.61 7.36 9.40 10.74
Ether extract (% ) 2.52 1.85 5.0 12.6
NFE(%) 44.8 45.9 41.8 34.9
Crude fibre (%) 21.0 24.1 24.5 23.0
Phosphorus (%) 0.13 0.16 0.15
Calcium (%) 1.12 1.14 1.13
Potassium (%) 1.60 1.22 2.14
increased by addition of sodium hydroxide and further improved by storing

the treated hay for up to three days (Myung et al., 1986).
The beneficial effect of alkali treatment was confirmed by Abou-Raya et
at. (1971) by feeding groundnut hay to adult sheep. They found that the
digestibility of all components, except N-free extract, was increased by
treatment with calcium hydroxide, sodium hydroxide, and sodium hydrox-
ide plus urea. However, the total dry matter was reduced by the hydroxide
treatments.
Chopping or grinding had no effect on the voluntary DM consumption
and digestibility coefficients of crude nutrients of groundnut haulms fed to
goats (Ayoade and Njewa, 1983/1984). Even so, in South Africa the
haulms are usually hammer-milled before being used in the ration (C.l.
Swanevelder, personal communication).
15.2.5 Haulm yields

Haulm yields vary markedly and are affected by variety, pre-harvest
conditions, disease level, time of digging, conditions after digging and
method of retrieval and storage.
Maximum dry matter yield of groundnut tops is reached before the
optimum lifting time for kernel yield (Wright et al., 1991). Under dryland
conditions, vegetative dry matter yield peaked at 90 days after planting
and, for most cultivars, declined steadily to digging at 130 days. Under
irrigation, this decline did not begin until 100-130 days after planting, with
different varieties showing different responses. Halevy and Hartzook
(1988) showed that the shedding of leaves and stems caused a reduction of
1386 kg from a potential haulm yield of 6006 kg/ha. This decline was also
recorded by Bell (1986), who observed the decline was accelerated by
foliar diseases.
Powell (1986), working with spanish varieties in central Nigeria, noted a
highly significant positive correlation between pod yield and yield of
vegetative dry matter. In the USA, spanish types yield 1-1.5 tonnes of hay
per tonne of pods, while runner types yield 1.5-2 tonnes per tonne of pods
(York and Colwell, 1951). This compares with results from southern
Queensland, Australia, where haulm:pod ratios are generally in the range
of 1 to 2: 1 for virginia bunch under dryland conditions and 0.6 to 1.2: 1 with
irrigation (G.R. Harch, personal communication). Ratios for spanish
varieties in the same dryland environment range from 1.4 to 1.7:1 and for
irrigated crops from 0.7 to 1:1.
15.2.6 Animal performance

Groundnut haulms have been fed to a range of animals including cattle
(Shukla et al., 1985), goats (Jamarun, 1984), pigs, rabbits (Aduku et al.,
1986), sheep and fish (Liao Boshu, personal communication), with varying
success in terms of animal performance and economic returns.
Goats fed groundnut haulms (6.5% digestible protein) gained weight
60% faster than on Digitaria hay (3.5% digestible protein) but at almost
double the cost per unit weight gain. Up to 75% groundnut hay can be
substituted for fresh ipil-ipil (Leucaena leucocephala) as a feed for goats
without reducing growth rates (Robles and Capitan, 1985). Similarly
groundnut haulms, when replacing maize stover, can almost double the
growth rate and increase the feed conversion efficiency of stall fed bul-
locks, although this advantage was lost through feeding ad lib. maize bran
with either feed (Addy and Thomas, 1977).
Groundnut straw can comprise 60% of the ration of growing lambs
without adversely affecting their growth rate and feed efficiency (Krishna
Mohan et al., 1985). However, as groundnut straw increases in the ration,
the cost per unit weight gain increases and the digestibility of all nutrients
tends to decrease. The most economical ration for growing lambs is 40%
groundnut hay and 60% protein concentrate (Durga Prasad et al., 1986).
Increasing the protein content of the concentrate can increase the intake of
groundnut hay by calves (Ahmed and Pollett, 1979).
15.2.7 Haymaking
Mechanical haymaking is feasible with groundnut herbage but losses are
high (Combellas et al., 1972). Groundnut hay in Australia is baled into
small rectangular bales weighing around 25 kg or larger round bales of
300-500 kg. The cost of baling groundnut haulms is higher than for other
hay because soil contamination leads to greater machinery maintenance
costs. Soil adhering to roots can make up 10% of the weight of groundnut
residues (Powell, 1986). In South Africa, sand contamination of baled
haulms can have a detrimental effect on animals' teeth (CJ. Swanevelder,
personal communication).
15.2.8 Storage
Long-term storage of groundnut hay in Australia is difficult. Mice are
attracted by unthreshed kernels and destroy hay bales that have been put
in sheds to reduce weather damage.
15.2.9 Aflatoxin
Aflatoxin poisoning may result from feeding groundnut haulms. Kernels
and shells present the greatest risk. Poisoning can result from animals
consuming ground nuts in the field, eating hay containing nuts or being fed
shells remaining after the kernels have been removed. Hay made from
groundnuts severely affected by drought, where threshing is uneconomic,
may have more nuts attached and thus present a higher risk, particularly
for young or weak animals with little or no other feed source.
Calves have been killed from eating groundnut hay containing pods with
high aflatoxin levels (McKenzie et ai., 1981). Jaundice, photosensitization,
diarrhoea, anorexia and depression were observed prior to death, while
post mortems showed haemorrhage, hepatocyte damage and bile duct
proliferation.
15.2.10 Chemical residues

Agricultural chemicals can improve the yield and quality of groundnut
haulms but their misuse can lead to haulm contamination, rendering them
unfit for use as animal fodder. Farmers in South Africa will even sacrifice a
little pod yield to ensure a good forage free from chemical residue (CJ.
Swanevelder, personal communication).
Haulm quality will generally be much higher from groundnuts treated
with a fungicide. By controlling leaf diseases, the leaf-to-stem ratio of the
haulms will be higher. However, Australian label requirements of some
fungicides (e.g. chlorothalonil) state that groundnut haulms should not be
fed to livestock. Other fungicides have a withholding period after spraying
before haulms can be fed; for example, groundnuts treated with cyproco-
nazole should not be grazed or cut for stock food for 14 days after
application. Sulphur-treated ground nuts have no withholding period.
Livestock should not be fed haulms and hulls from crops treated with
some nematicides, such as aldicarb, fenamiphos and ethoprophos.
Similarly the haulms from crops treated with herbicides, such as acifluor-
fen, sethoxydim and 2,4-DB, should not be fed to livestock in Virginia,
USA, yet haulms from crops sprayed with 2,4-DB in Queensland,
Australia, can be fed to livestock seven days after spraying.
TABLE 15.3 Nutrients removed from the soil by both groundnuts and haulms
(after Collins and Morris, 1913)
Nutrient content (kg)
Part of the plant Yield (kglha) N P K Ca Mg
Haulms 4540 89.8 5.2 77.8 44.5 17.2

Nuts 2250 69.4 6.7 19.7 3.2 3.5
Percent of nutrients
in hay 56.4 43.6 79.8 93.4 83.2
Organochlorine pesticides (DDT, dieldrin, endrin and BRC) have been

banned for crop use in many countries because of their long residual life in
soil and subsequent contamination of crops. To reduce such a risk in
Australia, soils in districts where organochlorines have been used are
tested prior to growing groundnuts.
15.2.11 Soil depletion

Removal of groundnut haulms affects the soil in two ways: firstly the direct
removal of nutrients from the soil, and secondly the depletion of organic
matter which is essential to maintain soil structure. Large quantities of
nitrogen and potassium are removed when haulms, as well as pods, are
taken from the soil (Collins and Morris, 1941) (Table 15.3).
In assessing the monetary value of groundnut hay the cost of replacing
these nutrients must be taken into account. At current prices in Australia,
to replace major and minor nutrients (e.g. N, P, K, Ca, Mg, S, Cu, B, Zn,
Mn) would cost approximately 35% of the market value of the hay.
A more difficult factor to measure is the long-term effect on organic
matter depletion. Most groundnut-growing soils are of a light texture and
organic matter is vital to maintain their structure and soil moisture-holding
capacity. Groundnuts grown in soils low in organic matter are more
susceptible to attack by Sclerotium rolfsii (K.J. Middleton, personal com-
munication) and diplodia blight (Diplodia natalensis) (Queensland
Department of Primary Industries, 1985).
Removing groundnut haulms may, therefore, provide a short-term gain
with serious long-term effects unless steps are taken to counter the effect of
nutrient and organic matter removal.
15.2.12 Disease reduction

Alternatively, removal of disease-affected parts of groundnut plants may
have an effect in reducing subsequent crop infection by lowering the level
of inoculum (K.J. Middleton, personal communication). Diseases that
Wild species 633
could be controlled partially in this way include leaf spots (Cercospora
arachidicola and Phaeoisariopsis personata) and verticillium wilt (Verticil-
lium dahliae).
15.3 WILD SPECIES
Resslar (1980) points out that many of the subgeneric epithets used in
current classifications are not valid according to the International Code of
Botanical Nomenclature (Stafleu, 1978). However, by virtue of common
usage, the classificatory system proposed by Gregory et al. (1973) and
modified by Krapovickas (1990) will be followed. Arachis wild species have
largely been collected to provide genetic material for groundnut improve-
ment programmes, particularly in relation to disease resistance (Stalker
and Moss, 1987). Simpson (1990) notes that some 1500 accessions of wild
species have been collected since the early 1950s, but less than half survive
in world collections. Krapovickas (1990) recognizes 77 wild species, and
most are represented by at least one accession in current collections
(Simpson, 1990). Only a small proportion of these has been widely evalu-
ated for forage potential.
One of the most valuable attributes displayed by many members of the
genus is persistence under grazing. Several have persisted in grazed pas-
tures for 30 to 50 years after planting, and others show clear indications of
being able to do so. Many of the tropical legumes released to date decline
over a period of 5 to 7 years, after which the pasture becomes unproduc-
tive. To date, most attention has been given to section Rhizomatosae, and
more recently to section Caulorhizae. However, a limited range of material
from other sections has been evaluated and shows considerable promise.
15.3.1 Rhizomatosae
(a) Description
Members of section Rhizomatosae are perennial and are characterized by
the presence of rhizomes. The section comprises two series:
Prorhizomatosae, a diploid group of somewhat delicate types, and
Eurhizomatosae, a more robust tetraploid group (Gregory et al., 1973)
containing A. glabrata and A. hagenbeckii, and two or three unnamed
species (Valls et al., 1985). Forage cultivars selected so far have been
drawn from Eurhizomatosae and are commonly referred to as rhizoma
peanuts in the USA.
The rhizomes, which form a dense tangled mat (mostly in the top 5 cm of
soil), vary in thickness from a few millimetres to about 1 cm in the sward of
a variety, with average thickness varying from variety to variety. They are
woody structures surrounded by a papery, brown, deciduous bark.
Contributing to the tangled effect of the rhizomes are the often heavy tap
Figure 15.1 Typical Eurhizomatosae growth habit, with horizontal rhizomes,

erect stems and vertical taproots (scale two-thirds actual size).
roots and a multitude of fine, fibrous roots, both of which bear the
characteristic small, oblate, aeschynomenoid nodules described by Corby
(1981). Herbaceous vegetative shoots arise from the rhizomes (Figure
15.1), producing a dense leafy sward in pure stands, which, when mature,
may vary from about 5 cm to 35 cm in depth, depending on variety. This
variation is partly attributable to growth habit, since in some varieties
stems tend to be decumbent and in others more erect, but also to inherent
differences in robustness.
Leaves are tetrafoliolate, with leaflets varying from accession to acces-
sion in overall leaflet shape from linear lanceolate to oblanceolate and
obovate, and in shape of leaflet apex from acute to obtuse mucronate
(Figure 15.2). Leaf size depends largely on plant vigour and frequency of
defoliation but also differs markedly among accessions. In their key in
Gregory et at. (1973), Krapovickas and Gregory describe flowers as 'large,
Wild species 635
Figure 15.2 Variety of leaf shapes in Eurhizomatosae.
standard (22-25 mm wide, 18-22 mm high), yellow, soft orange to brilliant

orange, no red veins on back of standard, wings usually not flared'.
However, flowers of some accessions may be somewhat smaller than
described, attaining a standard width of only 15 mm (E.G. Cook, unpub-
lished data). Krapovickas (1973) notes that it is difficult to find fruit of the
Rhizomatosae in the wild, although fruit can be set under cultivated
conditions. Personal observation is that some accessions fruit freely in an
immature state in cultivation but seedset declines rapidly with sward
development, despite profuse flowering. Niles (1990) ascribes this effect in
the American cultivar, Florigraze, to low pollen germination due to a dry
stigma and a high rate of embryo abortion due to competition in dense
canopies. The work of Lu et ai. (1990) suggests stigma morphology may
also be a contributing factor. Rhizomatous species studied had small
stigmas capable of carrying few pollen grains, in contrast with the large
stigmas of annual species which could accommodate more than 15 pollen
grains.
(b) Origin
Current theory holds that Arachis evolution follows movement and adap-
tation from an uplifted and dissected Tertiary peneplane in the region of
Mato Grosso in Brazil (Gregory et ai., 1980). With geocarpy a character-
istic of the genus, dispersal could only occur through vegetative advance,
peg elongation and erosive water movement. The other common dispersal
mechanism for legumes, voiding ingested seed, is unlikely since seed is
readily digested (Krapovickas, 1973). It is therefore not surprising that
geographical distribution of the various sections of Arachis largely follows
ancient and current drainage patterns. The Eurhizomatosae have been
collected between 13° and 28° S, in a triangular area extending south of
Corrientes (Argentina) to north of Rosario Oeste and east into Sao Paulo
state (both Brazil) (Gregory et ai., 1973). While they have mostly occurred
on red soils (Krapovickas, 1973), passport data in Gregory et al. (1973)
show that Eurhizomatosae have also been found on soils ranging from
light-coloured sands to fertile clays. Of the 186 accessions collected be-
tween 1936 and 1983, only 110 are currently conserved at the main germ-
plasm centres (Valls et ai., 1985).
(c) Cultivars
Rhizomatous cultivars have been released only in Florida. The first, Arb,
was registered by the Soil Conservation Service in Arcadia, Florida, fol-
lowing an evaluation programme commencing in 1937 (Prine, 1964). A
variety of A. giabrata, it was initially collected from 'a black rich earth in
the streets of Campo Grande, Mato Grosso do SuI' in Brazil (20° 27 ' S,
54° 37' W), and catalogued as PI 118457. The next cultivar, Arblick, was a
lower growing and more rapidly spreading type of A. glabrata, derived
from PI 262839. This accession was collected about 300 km south-west of
Campo Grande, near Bela Vista in Paraguay (22° 8' S, 56° 16' W). The
release of Florigraze by the University of Florida Institute of Food and
Agricultural Sciences and the USDA-SCS gave greater impetus to the
development of forage groundnuts in the USA. This variety arose as a
seedling on the border between year-old plots of the morphologically
distinct Arb and PI 151982, on the Agronomy Farm at Gainesville (Prine et
al., 1981). The new variety, initially catalogued GS-1, differed from the
other two in being fast spreading. It had more shoots per unit of soil
surface and a greater number of finer rhizomes than Arb - characteristics
more in common with PI 151982. Leaf shape was intermediate. In 1985,
IF AS and SCS released a further cultivar, Arbrook, a more robust variety
with coarser rhizomes than Florigraze, to improve the drought tolerance of
the species (Prine et al., 1990). Arbrook, introduced as PI 262817, was
collected on the top of a hill near Trinidad in Paraguay (27° 10' S,
55°40' W).
Research in eastern Australia has shown that another Paraguayan
Wild species 637
variety, GKP 9618 (introduced into Australia as CPI 93469), an
Argentinian variety, PI 231318/CPI 93483 and a third variety, PI 151982/
CPI 22762, whose origins are unknown, have sufficient promise to warrant
release in the near future. The first was collected near the Rio Peribebuy
(25° 12' S, 57° 5' W) (Gregory et al., 1973) but precise origins of the second
are not available (Anon., 1966).
(d) Adaptation
Forage research on Eurhizomatosae has been carried out in various South
American countries, USA, Australia, south-east Asia and India. Results to
date have shown various members of the series to have potential over a
latitudinal range from near the equator in Sulawesi (W.W. SHir, personal
communication) to 30° N in Louisiana (Caldwell et al., 1991) and 30° S in
New South Wales (G.P.M. Wilson, personal communication). Soil tex-
tures at evaluation sites range from the sands of Florida to the clay soils of
India and Australia. Soil fertility also varies markedly. The main prerequi-
site appears to be good drainage. Prine et al. (1981) make the point that
Florigraze 'is adapted to well-drained soils, but not to "flatwoods soils" or
to any soil which is subject to high water tables.' Experience in Queensland
supports this statement and shows that, although the suite of rhizomatous
accessions can survive temporary waterlogging, most vigorous stands are
obtained on well-drained flats and those parts of the slope not subject to
seepage. At the other extreme, various members of Eurhizomatosae have
demonstrated a strong degree of drought tolerance, one particular set
surviving under non-irrigated conditions at the ICRISA T Center near
Hyderabad, India, where the average annual rainfall of 750 mm is largely
concentrated over a 4-month period. PI 163452 has persisted for some
years in the Burnett region of southern Queensland (Jones and Rees,
1972), while Florigraze, not noted for its drought tolerance (Prine et al.,
1990), established and proved productive in semi-arid southern Texas
(Ocumpaugh, 1990) - both areas with similar annual rainfall totals to those
of Hyderabad but better distributed. The more drought tolerant Arbrook
is often more productive than Florigraze during dry periods, although
yields are similar under good rainfall conditions (Prine et al., 1990). In
severe drought, Prine et al. (1981) observed in Florigraze that top growth
may die off completely, the stand regenerating from rhizomes with the
onset of moist conditions. Vos and Jones (1986), assessing the importance
of rhizomes and stolons in legume persistence under grazing and using A.
glabrata PI 163452 and Lotus pedunculatus cv. Grasslands Maku as test
varieties, measured a small increase in Arachis rhizome yield per unit area
between an average rainfall year and an extremely dry year. In the more
moisture sensitive Lotus, rhizome yield was reduced by 96% .
A. glabrata and Eurhizomatosae generally are adapted to a wide range of
soil reaction. For Florigraze, Prine et al. (1981) suggest a soil pH of
5.8-6.5, although good yields have been obtained at pH 7.4 (Ocumpaugh,
1990), and no response to lime was obtained in an unidentified variety of
A. glabrata in any of three Florida soils with pH (in water) as low as 4.5
(Martinez and Blue, 1978). Niles et al. (1990), working with soils over a pH
range of 5.2-7.9, found that Florigraze showed a tendency to establish
better in soils with lower pH and higher aluminium. Contrasting somewhat
with this finding, Caldwell et al. (1991) obtained a response to dolomite in
Florigraze on a soil of pH 5.7. This is an enigmatic result, in that there was
no response to the addition of 2.5 t dolomite/ha producing a pH of 6.2, but
there was a response to 5 t dolomite/ha which raised the pH to 6.7. In
Australia, A. glabrata PI 163452/CPI 12121, perhaps the most widely
planted accession of the series, has been grown successfully on soils of
widely divergent reaction. This variety, along with several others, has
shown potential in a series of evaluation experiments in the wet tropics of
Malaysia and Indonesia on soils of pH 4.5-7 (W.W. Stilr, personal com-
munication). Results from more recent plantings of a wider range of
Eurhizomatosae on a variety of soil types in Australia suggest that the
adaptability to soil reaction displayed by selected members of the series is
common in the series. However, Reed and Ocumpaugh (1991) have
demonstrated variable manifestation of high pH-induced iron deficiency
symptoms among a range of rhizomatous accessions.
Since members of this series form an effective association with
Bradyrhizobium in the root nodules, nitrogen fertility of the soil is not
important to their long-term productivity. Nitrogen fertilizer applied to
pure swards of Florigraze had no effect on yield or persistence (Valentim et
al., 1988). However, when nitrogen was applied to a mixture of grass and
Florigraze, legume yields declined linearly with increasing nitrogen appli-
cation as grass yields increased (Valentim et al., 1986, 1988). It has also
been shown that nitrogen fertilizer applied at planting retards establish-
ment, probably due to stimulated weed competition and depressed peanut
nodulation (Adjei and Prine, 1976).
Perhaps the most important nutrient affecting pasture productivity, after
nitrogen, is phosphorus. Niles et al. (1990) were unable to demonstrate an
establishment response in Florigraze to applied phosphorus at 12 sites in
Florida soils with extractable phosphorus levels (Mehlich, 1953) ranging
from 11 to 317 ppm. It is worth noting that applications of potassium,
magnesium and sulphur also produced no measurable response. In pot
experiments, Blue et al. (1989), working on a Cameroon soil with
Florigraze as the test variety, and G.E. Rayment (personal communi-
cation), working on a dark clay loam from south-east Queensland with
GKP 9618/CPI 93469 as the test variety, demonstrated little or no response
in either variety to applied phosphorus. The Cameroon soil had a Mehlich
P level of 4.4 ppm and the Queensland soil a bicarbonate extractable P
level (Colwell, 1963) of 12 ppm. In both cases, test varieties from other
genera had proved responsive. From these experiments, and in view of the
Wild species 639
long-term persistence of a range of rhizomatous varieties at discontinued
unfertilized evaluation sites in southern Queensland, it appears that the
Eurhizomatosae share the characteristic common in the closely related
genus, Stylosanthes (Krapovickas, 1973) of being tolerant of low phos-
phorus levels in soils.
Eurhizomatosae have been successfully cultivated from about 0° to 30°
latitudinal range, although low temperatures in the higher latitudes can
cause significant damage. This may vary from death of plant tops, which
occurs even with light frosts, through to rhizome mortality under more
extreme conditions. Prine (1983) suggests rhizoma peanuts should be
grown in areas where temperatures do not fall below -10 0c. There appear
to be varietal differences in cold tolerance: Florigraze survived low tem-
peratures better than Arbrook (Prine et al. 1990); Arbrook was damaged
severely by freezes of lower than -12°C at Jay, Florida, and Americus,
Georgia (Prine et al., 1990); while Florigraze in Louisana, although mark-
edly reduced by a December temperature of -15°C had recovered to
about 70% coverage by November the following year. Such temperatures
are not experienced in those parts of eastern Australia where this series is
being evaluated.
(e) Yield
Rhizoma peanuts are best adapted to warm environments, preferring a
climate where temperatures are greater than 16°C for 5 months or longer
(Prine 1983). In southern Queensland, where monthly daily means range
from 14-15°C during June-August to about 25°C from December to
February, the average 6-weekly dry matter yield of 7 rhizomatous varieties
was negligible in the cool season but over 2 tlha in the warm season. In the
transitional period, yields of 0.1-l.0 t/ha were recorded (B.G. Cook,
unpublished). Likewise, dry matter yield/ha of Florigraze declined from
5.4 t in the summer to 2.9 t in the fall in Florida (Romero, 1986). This
apparent lower growth rate may be attributable to a partitioning shift from
shoots to rhizomes when daylength falls to 11 hours and temperatures to
20-22°C as demonstrated in Florigraze by Niles (1990). In the southern
Queensland experiment, the highest dry matter yield for a growing season
was 12 t/ha measured over six harvests in PI 231318/CPI 93483, and the
lowest was 3 t/ha in PI 338316/CPI 93464. Similar ranges in comparable
environments are reported for a multivarietal set in Florida by Prine
(1973), and for Florigraze in Louisiana (Caldwell et al., 1991) and in a
semi-arid Texas environment (Ocumpaugh 1990). Both Florigraze and
Arbrook have produced yields in excess of 16 t/ha in Florida (Prine et al.,
1986). Maximum yields obtained in the wet tropics of Malaysia and
Indonesia in a limited range of material were mostly below 5 t/ha/year
(W.W. SUir, personal communication), although these may have been
influenced by the shading effects of rubber trees and coconut palms.
However, comparisons between experiments should be viewed with
some caution, since yields of a particular variety may also be affected by
cutting height and frequency. Beltranena et al. (1981) obtained a 75%
increase in yield of Florigraze by taking 4 x 6-week cuts rather than
12 x 2-week cuts, and a 17% increase through cutting at 3.8 cm rather than
7.6 cm. Ortega et al (1992) measured similar responses in grazed mixed
swards of Florigraze and bermuda grass, where grazing intensity varied
from 500 to 2500 kg/ha of residual dry matter and grazing frequency from 7
to 63 days. They concluded that to maintain 80%, groundnut combinations
of at least 42 days between grazings and 1700 kg/ha residual dry matter, or
21 days between grazings and 2300 kg/ha residual dry matter, are
necessary.
In areas with a pronounced cool season, annual forage yields may be
improved by sowing cool-season species into the groundnut stand.
Dunavin (1990), working in Florida, obtained yields of 3-4.5 t DM/ha from
broadcasting seed of rye (Secale cereale) or italian ryegrass (Lotium multi-
florum) into a 5 cm stubble of Florigraze. Yields of Florigraze in the
following warm season were not affected. Cultivation to introduce the
winter crop of rye grass or oats (A vena sativa) in southern Queensland had
no detrimental effect on the ground nut stand, and actually enhanced its
spread.
(f) Feeding value

Eurhizomatosae produce unusually high quality feed for tropical pasture
legumes, often approaching the high levels found in temperate species such
as white clover (Trifolium repens) and lucerne (Medicago sativa). The most
commonly published estimates of forage quality are nitrogen percentage or
crude protein (N% x 6.25), in vitro organic matter digestibility (IVOMD)
and mineral levels, particularly phosphorus. In plants of Arb, Arblick and
Florigraze cut twice per season over 5 seasons, Prine (1973) measured N
levels ranging from 1.6% to 2.86%, P levels from 0.24% to 0.38%, and
IVOMD from 45.3% to 67.7%. With different accessions in south-eastern
Queensland, and plants cut at ground level every 6 weeks, N levels in the
dry matter ranged from 2.39%, to 3.56%, P levels from 0.3% to 0.54% and
IVOMD values from 69.9% to 77.7% (B.G. Cook, unpublished data).
Like yield, feed quality is influenced by height and frequency of cut.
Beltranena et al. (1981) increased leaf content from 72% to 93%, N levels
in the dry matter from 2.35% to 3.50%, and IVOMD from 64.8% to
74.3% by increasing cutting frequency from every 12 weeks to every 2
weeks. This improvement in quality is undoubtedly related to the higher
crude protein and IVOMD in leaves than in stems (Romero et al., 1987;
Saldivar et aI., 1990). Ocumpaugh (1990) demonstrated that nutritive value
is also affected by rainfall through its effect on dry matter production
patterns, leafiness, nitrogen levels in leaf and stem, and stem IVOMD.
Wild species 641
All the above aspects should be considered in order to achieve maximum
benefit from the crop. Beltranena et al. (1981) suggest from their data that
the crop be grazed every 4 weeks, green chopped or dehydrated as hay,
leaf meal or pellets every 4-6 weeks, or cut for hay every 8 weeks, though
Ocumpaugh (1990) suggests this should be less frequent in a semi-arid
environment. These qualities of rhizoma peanut have been shown to
translate into animal product using various classes of grazing animal.
Sollenberger et al. (1987) at Gainesville, Florida, achieved an average daily
liveweight gain (LWG) of 0.98 kg/head in yearling steers grazing Florigraze
compared with 0.37 kg/head in animals grazing Pensacola bahiagrass (Pas-
palum notatum) , reflecting the marked difference in quality between the
two forages. A similar pattern emerged at Brooksville (also in Florida)
where steers on Florigraze/grass pasture gained 0.68 and 0.90 kg/head in
consecutive years, compared with 0.52 and 0.50 kg/head on grass alone
(Williams et al., 1991). Although stocking rate in the first study was 43%
higher in the Pensacola than in Florigraze treatments, due to higher plant
growth rates, it is worth noting that L WG/ha was still 27% greater in the
ground nut than in the grass treatment. While it is common for L WG/head
to be greater on legume compared with grass pastures, it is unusual for
LWG/ha to be superior. However, rhizoma peanut compares favourably
with the 'king of legumes', lucerne or alfalfa (Medicago sativa). Gelaye et
al. (1990), feeding Florigraze and lucerne hay ad lib. to 4-5-month-old
20 kg goats, achieved daily L WGs of 63.8 g/head from the groundnut and
46.2 g/head from the lucerne. Feeds had similar analyses and were eaten in
similar quantities. This favourable comparison was borne out by Romero's
(1986) work with dairy cows, which gave similar milk yields from both
legumes, despite more favourable analysis of the lucerne due to its being
cut at an earlier stage than the groundnut. Romero et al. (1987) interpret
this in terms of slower ruminal breakdown of the protein in the groundnut
allowing a greater proportion of original protein to be utilized beyond the
rumen.
(g) Establishment
Since Eurhizomatosae generally produce poor seed yields, they are mostly
propagated from rhizomes. A commercial procedure has been developed
in Florida whereby rhizomes which have been produced in sandy soils are
lifted using a bermudagrass sprig harvester or a modified potato digger
(Prine et al., 1981). Where the potato digger is used, the rhizome mat is cut
into squares or rectangles with gangs of rolling coulters before lifting.
These pieces, normally about 30 cm square, can be planted in mounds no
more than 1.8 m apart, or preferably divided into four and planted 0.9 m
apart (Adjei and Prine, 1976). The recommended planting rate for both
Florigraze and Arbrook is at least 3.5 m 3 of rhizomes/ha (40 bushels/acre)
(Prine et al., 1981, 1986). Doubling this planting rate led to a first year
increment of 59% in one experiment involving Florigraze, but by the
second year differences were not significant (Valentim et at., 1988). A
similar trend was observed by Canudas et al. (1989) using lower planting
rates for Florigraze and Arbrook. The easiest way to establish rhizoma
peanut is to broadcast the rhizome pieces over the surface and cover them
using a disc harrow, then roll the soil to achieve rhizome/soil contact and to
retain moisture (Prine et at., 1986). Alternatively the rhizome pieces can be
planted in furrows from 3 cm deep in clay soils to 6.5 cm deep in coarse
sands. As a precaution, Prine et at. (1981) recommend inoculation of
rhizomes with an effective Bradyrhizobium strain at planting. However,
experience in Australia suggests that this is rarely necessary, due to the
presence of effective native bacteria either in the soil or already on the
rhizome surface.
In Florida, the preferred planting time is January or February, though
this could be as early as mid-December or as late as mid-March, allowing
for a slightly greater risk factor. At this stage, rhizomes are dormant. In
Australia, the slightly later southern hemisphere equivalent of August to
October has given good results, with January or February plantings result-
ing in very poor establishment. Saldivar et ai. (1992a) showed that levels of
nitrogen and total non-structural carbohydrate in the rhizomes were
highest in the coolest part of the year and that levels of both were reduced
by defoliation. These data support the preferred winter/spring planting
time and suggest that rhizome nurseries should not be defoliated until
immediately before planting. Defoliation also reduced rhizome production
(Saldivar et ai., 1992b).
Effective weed control is important during the relatively slow establish-
ment period. Canudas et ai. (1989) demonstrated this for both Florigraze
and Arbrook, achieving about double their ground cover and yield at the
end of the establishment year through controlling grasses and broadleaf
weeds. The broad-spectrum tolerance of rhizoma groundnut to herbicide
has facilitated preparation of a weed control programme incorporating the
use of pre-emergence applications of benefin, trifluralin or vernolate, post-
emergence applications of alachlor and dinoseb, and routine applications
of bentazon and 2,4-DB for broadleaf weed control, and sethoxydim
and fluazifopbutyl for grass control as required (Prine et ai., 1986).
Sethoxydim, but not dalapon, can be used to kill bermudagrass (Cynodon
dactyion) rhizomes contaminating groundnut rhizome planting material,
thus improving initial performance significantly (Canudas-Lara et at.,
1984). Mowing just above groundnut top growth may also reduce weed
competition and enhance establishment (Prine et al., 1986).
(h) Companion species

Although it might be important to suppress or control grasses during
establishment, it may well be desirable in the interest of long-term stability
Wild species 643
to introduce a grass into the groundnut sward to absorb nitrogen from the
legume system, thus reducing the likelihood of invasion by unpalatable
nitrophilous weed species. Valentim et al. (1986, 1988), using Florigraze in
association with a range of grasses, have demonstrated that, although
groundnut yields might be reduced due to the presence of a companion
grass, total pasture yields are at least the same and possibly enhanced.
By virtue of their unusual growth habit, Eurhizomatosae have been able
to grow effectively with grass species whose compact growth habit and
competitive ability may have precluded a successful association with other
legumes. Perhaps the most notorious of these grasses is Pensacola bahia-
grass (Paspalum notatum) although pangolagrass (Digitaria eriantha) and
bermudagrass are also recognized as being extremely competitive. Prine
(1964) records that Arb, once established, comprised 44%, 49% and 73%
of the top growth when grown in association with these respective grasses.
Prine et al. (1981) estimated that Florigraze, Arb and Arblick comprised,
on average over five seasons, 90%, 60% and 60% of the total forage yield
when grown with Pensacola bahiagrass. Valentim et al. (1986) found that
Florigraze comprised 86% of the forage yield in association with
Pensacola, 73% with bermudagrass and 81 % with Survenola digitgrass
(Digitaria x umfolozi) in the first year after establishment. In a later
experiment (Valentim et al., 1988), this time with Mott dwarf elephant
grass (Pennisetum purpureum) , Florigraze comprised 53% and 54% of the
forage yield in the first two years. All comparisons were in the absence of
added nitrogen. In Australia, the various accessions tested have been
grown successfully with similar grasses as well as with other sward-forming
species, rhodes grass (Chloris gayana) , paspalum or dallis grass (Paspalum
dilatatum) , brunswickgrass (P. nicorae) narrowleaf carpet grass (Axonopus
affinis) and blue couch grass (Digitaria didactyla) and tussock grasses such
as setaria (Setaria sphacelata var sericea) and the native black speargrass
(Heteropogon contortus). While there are many instances of long-term
persistence of rhizoma peanut in grass swards, it should be noted that at
Jay, in north-western Florida, the proportion of Florigraze in association
with a range of sward-forming grasses declined in the latter part of an
8-year period (Dunavin, 1992).
The ability to compete with grass associates varies from accession to
accession. In an experiment near Gympie in south-east Queensland com-
paring spread over 5 years of 31 Eurhizomatosae accessions through an
established grass sward under grazing, radial spread varied from a total of
3 cm in PI 338284/CPI 93473 to 143 cm in PI 262814/CPI 93475 (B.G.
Cook, unpublished). The American cultivars, Arb and Arblick, performed
poorly. Rate of spread in the absence of grass was much greater.
Calculating from the data of Adjei and Prine (1976), Florigraze spread at
2 m per year. A similar rate was observed with PI 338316/CPI 93464 in
south-east Queensland.
Competitive or spreading ability is not necessarily related to yielding
ability. In the Gympie study, those accessions with the better spreading
ability tended to be low growing, and those that had produced high dry
matter yields in other experiments performed relatively poorly in the
spread experiment. The exception was PI 231318/CPI 93483 which pro-
duced the highest yield and spread through grass at 14 cm per year, ranking
8 for rate of spread. The implication is that there is sufficient diversity
within the series to select hay types which are taller varieties mostly grown
in pure stands, pasture types which are capable of reasonable production
when grown with grasses, and the lower growing fast-spreading types
which may be more appropriate for soil conservation and amenity use.
(i) Alternative uses

Eurhizomatosae have potential beyond the provision of hay and forage by
virtue of their strong perenniality and competitiveness, their tolerance of
mowing and of herbicides, associated with their dense generally low
growth habit, and mid to deep green foliage with periodic flushes of
brightly coloured flowers. Prine et al. (1981, 1986) suggest that the
American cultivars could be used as ornamental ground covers and for
stabilizing road verges and embankments. These cultivars, in descending
order of suitability, are Arblick, Florigraze, Arb and Arbrook. They also
note that rhizoma peanuts have been planted as cover crop between rows
of orchard trees and vines but the long-term implications of such a practice
have not been determined. Among the current Australian collection, thc
most suitable accessions for amenity use are CPI 19898, PI 151982/CPI
22762, PI 338316/CPI 93464, GKP 9618/CPI 93469, PI 262814/CPI 93475
and PI 2628411CPI 93476. These are dense low-growing types and most of
them spread rapidly.
(j) Pests and diseases

High levels of resistance to insect and disease attack have been identified in
the Eurhizomatosae, although such resistance to anyone organism is not
necessarily universal throughout the series. Herbert and Stalker (1981)
found that, out of 55 rhizomatous accessions screened, 39 displayed a high
level of resistance to peanut stunt virus. Prine et al. (1986) noted that there
was also resistance to peanut stripe and mosaic viruses within the series.
Varying resistance to corn earworm (Heliothis zea) has been demonstrated
by Campbell et al. (1982); and to this and other insects - tobacco thrips
(Frankliniella fusca) and potato leafhopper (Empoasca fabae) - by Stalker
and Campbell (1983). All members of the Australian collection of the
series are resistant to races of organisms causing early leaf spot (Cerco-
spora arachidicola), late leaf spot (Phaeoisariopsis personata) and peanut
rust (Puccinia arachidis) in Australia (K.J. Middleton, personal communi-
cation). Although immunity to rust appears common in the Rhizomatosae,
Wild species 645
susceptible accessions exist (Ramanatha Rao, 1987). Stands of a number of

cultivated Eurhizomatosae were reduced or eliminated by rust near
Brasilia (E.A. Pizarro, personal communication). White mould (Sclero-
tium rolfsii) and lepto spot (Leptosphaerulina brisianna) have been ident-
ified on Arbrook, and leaf spots caused by Phyliosticta and Stemphylium
on both Florigraze and Arbrook, but none has caused long-term or serious
damage (Prine et al., 1981, 1986). Cotton root rot caused by the fungus
Phymatotrichum omnivorum has produced temporary reduction of stands
of various accessions in southern Texas (W.R. Ocumpaugh, personal
communication) .
Root-knot nematode Meloidogyne sp. can weaken the legume com-
ponent in pasture, so that level of resistance to these nematodes is often an
important determinant of the utility value of various legumes when grown
in association with other crops. The American cultivars Florigraze and
Arbrook proved highly resistant to the root-knot nematodes, M. arenaria
race I, M. javanica and M. incognita races I and II (Baltensperger et ai,
1986). Resistance to M. arena ria (Nelson et al., 1989) and M. hapla
(Castillo et al., 1973) has been identified in other rhizomatous accessions.
15.3.2 Caulorhizae
(a) Description
Section Caulorhizae is characterized by the presence of stolons. While
many Arachis have prostrate stems and may appear to be stoloniferous by
the rooting effect of the fruiting pegs, only the Caulorhizae actually form
regular nodal root systems (Gregory et al. 1973). The original accessions of
the two species within the section, A. pintoi and A. repens (Figure 15.3), are
morphologically distinct. However, more recent collections tend to be
intermediate between the two, making it difficult to ascribe them to one or
the other species (Valls, 1983). Descriptions will therefore largely apply to
the original types. Both species are strongly perennial, developing roots
along the stolons on those nodes touching the ground, or those in the high
humidity of a dense canopy. These roots, fine at first, ultimately develop
into large woody taproot systems, 1-2 cm in diameter near the crown. Roots
carry large numbers of small aeschynomenoid nodules. In southern
Queensland, A. pintoi may form a canopy up to 20 cm deep in a pure sward,
while A. rep ens rarely exceeds 10 cm. These depths may be exceeded when
either is grown with a taller growing grass, when the groundnut stolons
adopt a more ascendant growth habit. A. repens tends to produce a denser,
more closely appressed stolon mat than does A. pintoi. Leaves of both
are tetrafoliolate, with distal (apical) leaflets obovate and the proximal
(basal) leaflets oblong-obovate. Leaflet size and proportion vary between
species. A. pintoi leaflets grow to about 45 mm long and 35 mm wide,
while A. repens leaflets rarely exceed about 30 x 15 mm. Leaflet
Figure 15.3 (a) Part of Arachis pintoi plant with stolon, taproot, foliage, flower
and pods; (b) variation in pod types of A. pintoi; (c) leaf of A. repens. (Scales: (a)
2/3 actual size, (b) 1.5 x actual size, (c) 1-1.5 x actual size.)
apices are mostly obtuse and bases slightly cordate in A. pintoi, the apices
of A. rep ens being mostly mucronate acute, and the bases cuneate to
obtuse. A. repens also lacks the prominent spreading tuberculate hairs that
are common on the proximal leaflet of A. pintoi.
Both species have relatively small yellow flowers, although one of the
recent collections which is closer to A. repens has orange flowers (Valls,
1983). One of the more significant differences between the two species is
that A. pintoi is capable of producing high yields of seed (Cook and
Franklin, 1988) and the original A. repens rarely produces seed at all. A.
pintoi pods are mostly single-seeded, about 6 mm in diameter and 11 mm
long, and borne usually singly, on pegs 1-27 cm long with the terminal pod
being prominently beaked. There are about 6000-8000 seed-in-pod/kg
(Oram, 1990).
Wild species 647
(b) Origin
A. pintoi was first collected in April 1954 by Professor Geraldo C.P. Pinto
near the mouth of the lequitinhonha River in Brazil at 15° 52' S, 39° 6' W,
and 50 m A.S.L. (Gregory et al., 1973). Valls (1983) and C.E. Simpson
(personal communication), after visiting the collection site and many sites
upriver, concluded that this material had been transported downriver from
around Ara~uai, that the collection site no longer exists due to southward
displacement of the river mouth, and that the altitude cited above would
more likely have been 5 m A.S.L.. It has travelled a circuitous route
around the world - from the collection site, in turn to Cruz das Almas
(Brazil), Corrientes (Argentina), Experiment (USA), Brisbane (Austra-
lia) and Cali (Colombia), ultimately returning to Brazil at Belem in the
north. It has now been catalogued under a variety of collection numbers
and accession numbers: GK 12787 = PI 338447 (US), HL 323 = PI 338314
= CPI 58113 (Australia) = CIAT 17434 (Colombia).
The site where Dr Thomas Dalton first discovered A. repens was at
600-700 m A.S.L. along the lequitai River (18° 10' S, 43° 26' W). Gregory
et al. (1980), believing this to be a tributary of the lequitinhonha rather
than of the Sao Francisco, recorded that Caulorhizae has been found only
in the valley of the lequitinhonha. Subsequent collecting expeditions have
extended the known limits of the section to include the valleys of the Sao
Francisco (Valls, 1983) and Tocantins (C.E. Simpson, personal communi-
cation). The original A. rep ens collection is now catalogued under seven
different collection numbers, seven USA accession numbers and a variety
of other accession numbers throughout the world.
A. pintoi has generally been collected in infertile red, sandy loam river-
bottom soils of high aluminium saturation, appearing to do best in low
areas which are wet to flooded during the wet season. Rainfall in the region
during the wet season (October to May) is about 2000 mm, a further
200 mm falling during the remainder of the year. The native vegetation is
mata baixo or low forest, with trees to 5 m and very little sunlight reaching
the forest floor (C.E. Simpson, personal communication).
A. repens originates from similar environments. The only published
noting about the first collection site was 'from an area sometimes inun-
dated by water' (Anon., 1967). One recent collection was from a grey, silty
loam soil, where the plants were growing among grasses in standing water,
and another from a red silty clay where plants were growing on the
roadside and in low areas (C.E. Simpson, personal communication).
(c) Cultivars
The only cultivar from Caulorhizae is A. pintoi PI 338314, which was
introduced into Australia as CPI 58113 in 1972, released commercially
in 1987 and registered as cv. Amarillo (mistakenly believing this to be
Portuguese for 'yellow', referring to the flower co.lour) in 1990 (Oram,
1990). Amarillo has been exported to Colombia, Indonesia, Malaysia,
Thailand, Sri Lanka and various Pacific countries including Fiji, Western
Samoa and Tonga.
(d) Adaptation
In general, A. pintoi and A. repens are well adapted to moderately
drained, or seasonally inundated or poorly drained soils in the humid
tropics and subtropics. They have performed well between latitudes of
1° 30' near Manado in North Sulawesi, Indonesia (Kaligis and Sumolang,
1991), to 29° 30' at Grafton in New South Wales, Australia (G.P.M.
Wilson, personal communication), in areas receiving average annual
rainfall from about 1100 mm to more than 3000 mm. Soil textures at
successful evaluation and development sites have ranged from sands to
well structured clays, and soil reaction mostly from very acid to neutral.
Due to more intensive testing of A. pintoi, there is a more detailed
understanding of its adaptational range than for A .. rep ens . The only soil
texture limitation for A. pintoi appears to be on poorly structured, medium
to heavy clays, often referred to as 'puggy' soils. In two instances in
southern Queensland, it has grown poorly or failed to persist on such
soils (R.M. Jones, B.G. Cook, unpublished); this does not appear to be
related to drainage, since experience in Brazil shows it to be well adapted
to seasonally waterlogged conditions (Anon., 1989b). It appears to be
less sensitive than many species to pH, having grown well on soils
with pH values ranging from 4.3 (in water) on the Llanos Orientales of
Colombia (Grof, 1985) to 7.2 (1:5 water) on the Darling Downs of south-
east Queensland (J.N. Coote, personal communication). As might be
expected, the adaptation to extreme acidity is accompanied by a tolerance
of soluble aluminium and manganese. G.R. Rayment (unpublished data)
demonstrated tolerance of manganese levels which would have been detri-
mental to many other legumes. Aluminium tolerance is indicated by soil
analysis data from Grot's (1985) experiment, which showed an 86%
aluminium saturation of the exchange complex.
A. pintoi may have a greater requirement for soil phosphorus than the
rhizomatous species (section 15.3.1( d)). In a pot trial comparison between
the rhizomatous GKP 9618/CPI 93469 and A. pintoi, using a soil of 12 ppm
available P (bicarbonate extract; Colwell, 1963), the former gave little or
no response, while the latter responded up to the equivalent of 50 kg/ha of
applied phosphorus. Dextre (1984), Turnour (1987) and J.X. Kui (unpub-
lished data) also obtained significant responses in A. pintoi to single rate
applications of 20, 100 and 40 kg/ha of phosphorus to soils of 2.7-4.9 ppm
(extractant not reported), 13 ppm and 19 ppm (both bicarbonate extract)
respectively.
While A. pintoi and A. repens are adapted over a wide latitudinal range,
Wild species 649
best performance is obtained in areas not subject to frost. Top growth is
'burnt' by frost but stands recover from unaffected stolons or crowns.
One of the most significant characteristics of A. pintoi is its high level of
shade tolerance. In a study by the University of Queensland comparing 84
legume accessions from many genera, A. pintoi was the most shade toler-
ant; at 20% light it produced 48% of the mean of yields produced at 100%
and 70% light (W.W. Stiir, 1991). This has far-reaching implications in its
use as a forage and ground cover. A. repens was not as shade tolerant as
A. pintoi; at 20% light it produced only 28% of the mean of yields
produced at 100% and 70% light.
(e) Yield
The magnitude of dry matter yields reported for a particular species is
often strongly dependent on height and frequency of cut. It is not surpris-
ing, then, to find that yields from experiments in various countries differ
markedly. In Colombia, harvesting every 4 weeks at 3 cm, Grof (1985)
obtained dry matter yields from 5.2 to 9.6 t/ha, depending on the com-
panion grass. Yields in Costa Rica varied from 2.6 t/ha (11 cuts/yr) at San
Isidro to 10.9 t/ha (9 cuts) at Guapiles (Anon., 1990b). Working in matur-
ing rubber plantation in Malaysia, Ng (1991) recorded only 2.4 t DM/ha.
Kaligis and Sumolang (1991) and Rika et al. (1991), working in coconut
plantations in north Sulawesi and Bali in Indonesia, obtained yields of 4.8
and 2.7 t DM/ha for A. pintoi and 4.4 and 3.2 t DM/ha for A. repens. In all
south-east Asian situations, Arachis was grown in pure stands and cut at 8-
weekly intervals. In the humid sub tropics of south-eastern Queensland,
harvesting pure stands at 2-3 cm every 6 weeks, irrigated and unirrigated
yields of 7.3 and 6.5 t DM/ha were obtained from cv. Amarillo (B.G.
Cook, unpublished).
In the Guapiles experiment, two more recently collected accessions of
A. pintoi (ClAT 18744, 18747) produced similar yields to the original
collection (ClAT 17434) while others (CIAT 18745, 18746) produced only
3.7 and 3.8 t DM/ha.
(f) Feeding value

Both A. pintoi and A. repens are readily eaten by grazing livestock but
information on feeding value is restricted to the former. Animals may
select A. pintoi over other components of a mixed pasture. In pastures
comprising A. pintoi and different species of the grass genus, Brachiaria,
selection indices of 0.57 to 0.61 were calculated for three of the four
mixtures, the other being 0.48 (Lascano and Thomas, 1988). Selection
index relates the proportion of a species in the diet to the proportion in the
pasture, a level <0.5 indicating selection against and >0.5 selection in
favour (Chesson, 1983). Analysis of the feed on offer revealed leaf:stem
ratios of about 0.24 in the dry season and 0.97 in the wet season but, unlike
most forage legume species, there was little difference between IVOMD of
leaf and stem, the greatest difference being between IVOMD of leaf in the
dry (67%) and the wet season (60%). Mean value of crude protein (CP) in
the leaf was l3.9% in the dry season and 18.0% in the wet season, while
stem values were about 11 % in both seasons. In earlier measurements
from the same pastures, Grof (1985) measured average crude protein
levels of 14.8% in the first year and 16.6% in the second, and phosphorus
levels in the dry matter of 0.18-0.20%. With an average stocking rate of
2.4 animal units/ha (1 a.u. = 420 kg), these pastures proved capable of
producing 515 g/head/day over a 594-day grazing period. In south-east
Queensland, a small plot cutting trial produced dry matter of similar high
quality to that obtained in the above experiment in Colombia - 20% CP,
73% IVOMD and 0.35% P (B.G. Cook, unpublished data), the latter
figure probably reflecting the high levels of phosphatic fertilizer applied.
(g) Establishment
Due to poor seed set, A. repens is propagated only vegetatively. It strikes
readily from lengths of stolon with two to three nodes, providing the soil is
kept moist.
A. pintoi produces high seed yields and may therefore be propagated
from either seed or cuttings. A laboratory technique for in vitro culture has
also been described (Burtnik and Mroginski, 1985) but has no immediate
application in commercial pasture establishment. The simplest method is
to establish from seed. Freshly harvested seed should be dried at 35-40 °C
for 10 days to improve germination (1. Butler, unpublished data). Cook
and Franklin (1988) describe a method of precision planting to establish a
seed crop. Seed should be inoculated with a specific strain of
Bradyrhizobium prior to planting. Early evaluation of the species was
hampered by lack of a highly effective strain of root nodule bacteria, but
the strain CIAT 3101, initially isolated from Centrosema macrocarpum,
was selected after a screening programme in Colombia (Sylvester-Bradley
et ai., 1988). This strain, different from that used with A. hypogaea, is now
used in all commercial plantings. For the seed crop, a sowing rate of 25 kg
seed-in-pod/ha was used, but 10 kg/ha is adequate for pasture establish-
ment providing seed is of good quality. Seed is best sown 2-6 cm deep.
Specialized planting machinery is most economical of seed but other
methods are available. Perhaps the most practical method is to broadcast
seed onto a fairly well prepared seed-bed, cultivate to cover the seed, and
roll to achieve good soil/seed contact. Some coverage is essential as
surface-sown seed germinates poorly (Ogawa et ai., 1990) and is attractive
to birds and rodents (B.G. Cook, unpublished data). Ogawa et ai. (1990)
noted that some seeds that did germinate failed to develop because the
thick radicle could not penetrate the hard surface layer of uncultivated soil.
Wild species 651
A. pintoi is tolerant of a range of pre- and post-emergence herbicides
facilitating effective chemical weed control in an establishing stand. Its low
growth habit in the early stages permits use of rope-wick herbicide appli-
cators (Cook and Franklin, 1988). Sowing time depends on seasonal
constraints such as probability of frost in the higher latitudes, and onset of
wet and dry seasons in monsoonal climates. Seedlings establish quickly and
complete groundcover can be achieved in 5 months from the higher sowing
rate given above. Seed is not always available, due to difficulties in seed
production and storage, but established stands may be available as a source
of planting material. Asakawa and Ramirez (1989) describe an effective
method of manual or mechanical vegetative establishment used in
Colombia. They suggest planting 20 cm lengths of stolon about 15 cm deep
in rows 35 cm apart or, if planted with a grass, in rows 1 m apart. Cuttings
should be inoculated with Bradyrhizobium using an adhesive such as
molasses.
(h) Seed production

A. pintoi seedlings usually commence flowering 3-4 weeks after emerg-
ence. The flowering continues through much of the growing season, with
periodic flushes apparently in response to improved soil moisture con-
ditions (Cook and Franklin, 1988). In the sub tropics , flowering declines
with the onset of cooler conditions in May, but in the tropics it continues
through most of the year (D.G. Cooksley, personal communication).
Initial flowers usually produce few pegs, and pegs produced during dry
conditions fail to develop fruit. However, with these exceptions, seed set
spans at least 5 months in subtropical Queensland, where all Australian
seed is currently produced. By the time end-of-season fruit have matured,
pegs of early-set fruit have decayed, giving low seed yields using con-
ventional groundnut harvesting techniques. Cook and Franklin (1988)
describe a more efficient harvesting principle that involves sieving the
whole soil volume through rotating, coaxial, cylindrical screens once the
seed is mature. Seed crops are grown in sandy soil to facilitate separation
of seed and soil. This system, which has now been commercialized, has
produced yields of about 1 t/ha of seed-in-pod per year. It is necessary to
rotary-hoe prior to harvest but only a small proportion of seed is damaged
in the process. No yield response or reduction in the proportion of 'pops'
has been obtained through applications of calcium as gypsum or as lime in
experiments at Gympie (D.S. Loch, unpublished data). There is no need
to resow seed crops, since stands recover from unharvested seed and from
new crowns formed at depth on taproots severed by the rotary hoe. In
Costa Rica, Diulgheroff et al. (1990), also using a sieving technique,
obtained seed yields ranging from 0.6 to 2.0 t/ha from experimental plots.
Smallholder farmers at Yapacani, near Santa Cruz in Bolivia, have pro-
duced large amounts of seed, at times reaching 2.6 t/ha, using simple
cradle-mounted flat sieves and digging hoes (E. Ramirez, personal com-
munication). Some of the more recently collected A. pintoi varieties have
proved capable of seed yields of about 1.5 t/ha in Peru in an experiment in
which CIAT 17434 (the original variety) inexplicably produced no seed
(Anon., 1990a).
(i) Companion species

These stoloniferous Arachis species are capable of forming stable associ-
ations with a variety of grasses. In the native habitat, A. pintoi has been
collected growing with Panicum maximum, Eriochloa polystachia and
Paspalum sp. and A. repens with Paspalum and Cynodon dactylon (C.E.
Simpson, personal communication). In Colombia, Grof (1985) showed
that A. pintoi could also grow successfully with the competitive Brachiaria
species (dictyoneura, brizantha, humidicola and ruziziensis) comprising
20-45% of the pasture. In another wet tropical environment at South
Johnstone in north Queensland, A. pintoi has proved capable of forming a
balanced pasture with B. decumbens. In the subtropics, it has been grown
with a wide range of native and introduced species (Oram, 1990). A. repens
has shown similar persistence in a variety of grasses in tropical and sub-
tropical Queensland. In an experiment in northern Queensland, both A.
pintoi and A. rep ens have spread about 1.5 m in 15 months, when grass
competition was initially low (D.G. Cooksley, unpublished data). Spread
with vigorous grass competition has been slower in the subtropics. A.
pintoi spread at 5 cm/yr over 5 years, and A. repens at up to 15 cm/yr (B.G.
Cook, unpublished data). R.M. Jones (unpublished data) measured a
similar rate of spread in A. rep ens over 15 years at Beerwah in south-
eastern Queensland, under competition from Axonopus affinis and
Digitaria eriantha subsp. pentzii.
(j) Alternative uses

Both Caulorhizae species have application other than as forages, by virtue
of sward-forming habit, shade tolerance or periodic flushes of flowering.
Soil loss is a serious problem in orchards and plantations. Ground-cover is
often destroyed intentionally to minimize competition with the developing
crop, or to facilitate ease of movement through the crop, or it may simply
succumb to increasing shade from the developing crop. Once ground-cover
is lost, erosion becomes a problem. Dwyer et al. (1989) suggest A. pintoi as
a living mulch, that should not compete with the crop to any great extent
because of its low nutrient demand and its ability to fix its own nitrogen.
However, G. Johns (Anon., 1991) recommends caution with this practice
in bananas: despite an improvement in soil fertility under A. pintoi at
Alstonville in northern New South Wales, fruit production over 3 years
was reduced by about 13% compared with a crop on bare soil. He suggests
Wild species 653
that this may have been due to reduced soil temperatures rather than
competition. The benefit is that there is no soil loss, which in turn implies a
more sustainable practice.
A. pintoi is also being used in north Queensland for bank stabilization
and as an ornamental ground-cover in shaded or erosion-prone areas
(P. Bolton, personal communication). Although A. repens has not been
similarly exploited in Queensland, it is used as an ornamental in its native
environment. C.E. Simpson (personal communication) observed A. repens
growing in domestic and commercial gardens during a collection trip in
Brazil, where it is locally known as 'grama amendoim' or groundnut grass.
PI 338277 was used principally for lawn planting at the collection site
(Anon., 1970).
(k) Pests and diseases

Neither of the original collections of the two species is severely affected by
disease. Both appear to be resistant to the common groundnut diseases,
rust (Puccinia arachidis) and leaf spots (Cercospora arachidicola) and
Phaeoisariopsis personata) (K.J. Middleton, unpublished data), although
C.E. Simpson (personal communication) has observed late leaf spot on
leaves of one of the more recent A. repens collections. A. pintoi CIAT
17434 has proved susceptible to two fungal diseases of the commercial
peanut - peanut scab (Sphaceloma arachidis) in Colombia, and pepper leaf
spot (Leptosphaerulina arachidicola) in Costa Rica (Anon., 1989a,c) -
both having only a limited effect on productivity. Two other lines, CIAT
18752 and 18746, were resistant to Sphaceloma in a field screening. Other
fungi - Phomopsis sp., Cylindrocladium sp. and Colletotrichum gloeospor-
ioides - have been isolated from leaf spots on A. pintoi in Australia but
none is serious or widespread. A. pintoi and A. repens are both subject to
often extensive black stem lesions, which apparently have little effect on
the overall vigour of the plant. Colletotrichum gloeosporioides has been
isolated from these lesions in Australia (Oram, 1990) and C. truncatum (C.
dematium) in Colombia (Grof, 1985). Inoculation of A. pintoi with strains
of C. gloeosporioides that cause severe damage to stands of the related
genera, Stylosanthes and Aeschynomene, failed to produce infection (Vini-
jisanum et al., 1987), suggesting that these strains are different from those
mentioned above.
Although a small proportion of leaves of A. pintoi cv. Amarillo in
Australia are variegated, no virus particles have been observed in the sap
under electron microscopic examination. However, peanut mottle virus
has been isolated from material showing virus-like symptoms in Colombia
(Anon., 1990c). Resistance to peanut stunt virus has been recognized in
A. repens (Herbert and Stalker, 1981).
Nematodes and insects have not proved to be a field problem with either
species. A. pintoi has a high level of resistance to Meloidogyne incognita
and M. arenaria, and moderate resistance to M. hapla, but is susceptible to
root lesion nematode (Pratylenchus brachyurus) (R.W McLeod and P.e.
O'Brien, personal communication). Stalker and Campbell (1983) demon-
strated resistance to tobacco thrips, potato leafhopper and corn earworm
in A. repens.
Perhaps the greatest potential problem lies not with the usual micro-
organisms but with rodents. In several situations, rat and mouse popu-
lations have increased in response to build-up of seed under A. pintoi.
While this has had no great effect on the legume stand, it can have
undesirable consequences with the associated tree crop, through excessive
soil aeration and drying from the burrowing animals, and loss of fruit from
the trees as the animals seek an alternative diet.
15.3.3 Other species

Various other species have been identified as having forage potential.
Following a plant collecting trip in 1947-48 through parts of Argentina,
Uruguay, Paraguay and Brazil, Hartley (1949) nominated a number of
species he considered to have merit by virtue of performance under grazing
in their natural habitat. Unfortunately, as intimated by Gregory et al.
(1973), specific affiliations at that time are different from those that would
be applied today. Although Hartley provides reasonable descriptions of
material collected, in many cases it is insufficient to identify with any
degree of certainty to which current species he is alluding. His selection of
highest potential is clear: through reference to rhizomes, his A. prostrata is
obviously the current A. glabrata or, at least, section Rhizomatosae. The
identity of his second choice A. glabrata is somewhat clouded. It is poss-
ible, from the plant descriptions and the distributions given, that these
accessions and those he assigns to A. diogoi ('one of the most promising
forage species') mayall, or in part, belong to the sections Erectoides and
Procumbensae. Hartley's judgement has been borne out in that members
of these sections have come to prominence in Queensland and in Florida.
The Queensland accession is noteworthy by its record of persistence.
Now simply accessed as CQ 1780, this line probably derives from material
initially identified as A. prostrata CPI 6930 or A. diogoi CPI 8987-8 (Miles,
1949). These lines were planted in the late 1930s or early 1940s at
Fitzroyvale near Rockhampton in central Queensland. Despite extremely
heavy grazing and lack of fertilizer, one type has persisted and spread at
this site. This line has recently been evaluated at other sites in coastal and
sub coastal Queensland, and appears to be well adapted to subhumid
environments (B.G. Cook, unpublished data).
The variety showing early promise in Florida, PI 446898, was collected
by Professor A.E. Kretschmer, Jr, on the Pantanal of Brazil (20° 7' S,
56° 43' W), where it was growing in 10-20 cm of standing water, on a clay
soil of pH 6.0-7.0 (Kretschmer and Wilson, 1988). This accession has a
Breeding 655
number of desirable attributes including fair yields of high quality forage
(Kretschmer and Wilson, 1988; B.G. Cook, unpublished data), resistance
to early and late leaf spot and rust (K.J. Middleton, unpublished data) and
high seed yields (D.S. Loch, unpublished data). It has performed weIl
under grazing in Florida but has not persisted well under grazing in coastal
Queensland, possibly due to attack by various root-knot nematodes and
several fungal diseases. Further evaluation is required.
All material considered thus far has been perennial. Beaty et ai. (1968)
assessed the annual, A. monticola PI 263393, for its persistence in swards
of Pensacola bahia grass and Coastal bermudagrass. Results were encour-
aging but this accession, when screened for disease susceptibility in
Queensland, proved susceptible to rust (K.J. Middleton, unpublished
data). It is important that forage varieties are not susceptible to the more
serious peanut diseases, not only because of the disease effect on forage
productivity but also because diseased plant parts are a source of inoculum
for nearby commercial crops.
15.4 BREEDING
Considering that the genus Arachis possesses many desirable forage attri-
butes among its various sections and species, it is remarkable that so little
breeding work appears to have been carried out with the specific aim of
producing improved forage types. Gregory and Gregory (1979) demon-
strated intra- and intersectional compatibility from a programme involving
a total of 1075 crosses among 91 parental forms representing at least 50
species. Although the success rate was not high, the work does demon-
strate some potential for improvement through breeding. Such improve-
ment has already been achieved at Coimbatore, India, from a cross
between the diploid A. diogoi and tetraploid A. glabrata (Chandrasekhar,
1980). The hybrid, which it is suggested be called cultivar Forachis, pro-
duces higher yields than either parent, yet retains the desirable rhizoma-
tous growth habit of A. glabrata. Gregory and Gregory (1979) showed that
A. giabrata crossed successfully with series Annuae of section Arachis and
series Tetrafoiiolatae and Procumbensae of section Erectoides, but not with
series Perennes of section Arachis, of which A. diogoi is a member. This
places doubt on the identity of the non-rhizomatous parent of Forachis.
Natural hybrids are common in pasture nurseries where representatives
of several sections are grown in close proximity. The American cultivar
Florigraze arose as an intrasectional cross under such circumstances at
Gainesville, Florida (Prine et ai., 1981). A probable intrasectional cross
occurred at Grafton in New South Wales, where a type, intermediate
between A. repens and A. pintoi, appeared in a plot of A. pintoi growing
close to a plot of A. rep ens (G.P.M. Wilson, personal communication).
No other compatible types were growing nearby. Intersectional crosses
between Erectoidesl Procumbensae and Rhizomatosae have also developed
at Grafton, and at Gympie, in south-east Queensland. These have
appeared in plots of the freely seeding Erectoides accession, CQ 1780, and
Procumbensae accession, PI 446898. The hybrids do not flower; they
have different leaf characteristics from the female parent and produce a
multitude of shoots arising from short rhizomes. Similar types have
been observed at Gainesville, Florida (K.H. Quesenberry, personal com-
munication). None of these appears to have an agronomic advantage over
either parent. Other possible spontaneous intersectional hybrids have
been observed in Costa Rica, with A. pintoi (Cauiorhizae) as the female
parent and PI 446898 (Procumbensae) as the male (P. Argel, personal
communication) .
However, once the limitations of the wild types have been more com-
pletely identified, there seems real potential to incorporate valuable attri-
butes into more promising types - say nematode resistance into PI 446898-
or to produce a completely different type combining desirable qualities
from several sources, using the various breeding techniques available.
15.5 CONCLUSION
The genus Arachis originates mainly from Brazil but is also found in
Argentina, Bolivia, Paraguay and Uruguay, in an area extending from near
the Equator to 34° S, and from the Atlantic coast to the foothills of the
Andes. The altitude range is from sea level to 1450 m. Gregory et ai. (1980)
describe the diversity of habitat:
It is found among vegetation types from broken forest to the open
grasslands of 'Pantanal' and 'Campo', in 'Caatinga', 'Parque',
'Chaco', and 'Cerrado'. Its species grow submerged, among stones
bathed with water, in dry gravel, and in flood-plain alluvium. They
are found from semi-arid locations to regions that receive 2000 mm or
more of rain per year and in regions subject to great floods and
intense droughts. They grow most commonly in friable or somewhat
sandy soils, but also in soils that vary from almost pure sand to
reduced, humic clays and sands, various alluvia, dark red humic
latosols, to pure lateritic caps and gravel. Underlying parent rock
may be sandstone, limestone, basic gneiss, granite, basaltic lava, and
less often mica and mica schists.
There are annual, biennial and perennial types, with procumbent, decum-
bent or erect growth habits. Prostrate stems develop nodal roots in some
species; in other species, rhizomes develop. Plants may have simple tap-
roots or varying degrees of tuberous development. Seed is produced
underground on vertical or horizontal pegs. The genus has been divided
into sections based on these characters. To date, forage potential has been
recognized and commercialized in only three of these sections - Arachis,
References 657
Rhizomatosae and Caulorhizae. Work with material from these sections
has demonstrated a number of particularly desirable characteristics within
the genus, though not all necessarily possessed by anyone variety - high
yields and high quality of forage, high palatability, excellent haymaking
qualities, persistence under intensive grazing and among competitive
grasses, tolerance of low fertility and high aluminium and manganese,
good drought tolerance, and minimal losses due to pests and disease.
These characteristics have translated successfully to animal production.
Outside the native habitat, the bulk of forage derived from Arachis has
come from haulms from the cultivated groundnut, A. hypogaea. At this
stage, only a relatively small proportion of the world collection of wild
types has been evaluated for forage potential in the tropics and subtropics.
With the diversity of growth forms and adaptation within Arachis, there is
little doubt that, as more of the collection finds its way into forage
germ plasm banks, the contribution of the genus to world livestock pro-
duction will increase markedly.
ACKNOWLEDGEMENTS
The authors gratefully acknowledge the efforts of Mr Will Smith, botanical

illustrator, Botany Branch, QDPI, in preparing the splendid illustrations
used in this chapter.
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Rome. pp. 10-17.
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Stalker, H.T. and Moss. J.P. (1987) Speciation, cytogenetics. and utilization of Arachis
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Stalker, H.T. and Campbell, W. V. (1983) Resistance of wild species of peanut to an insect
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CHAPTER 16
The groundnut in farming

systems and the rural economy
- a global view
J. Smartt
It is perhaps axiomatic to state that the groundnut crop is produced in

farming systems ranging from the most primitive to the most advanced.
The role it performs in these systems varies considerably, from labour-
intensive shifting cultivation to highly capital-intensive systems in North
America and Australia.
The groundnut has an enormous advantage vis-a-vis many other crops in
economies at or above subsistence level in that it has a dual-purpose role: it
is both a subsistence and a cash crop, in contrast to beverage and industrial
crops. Although it is a highly input-intensive crop, the nature of the inputs
can be varied according to local circumstances; it is energy demanding and
how these demands are met is open to alternatives. In areas where labour
is abundant, all operations can be carried out manually; where it is in short
supply, most can be mechanized. In intermediate situations it is possible,
by appropriate injection of capital and energy resources, to optimize
production efficiency for any particular set of circumstances. The develop-
ment of intermediate technologies is an important issue in developing
(as against stagnant) agricultural economies.
16.1 FARMING SYSTEMS
To consider the varied roles that ground nuts can perform, it is appropriate
to appreciate the actual diversity of farming systems in which it can be a
component. Farming systems can be classified according to a number of
different criteria. Ruthenberg (1980) classifies them on the following bases:
1. Type of land management
2. Intensity of cropping
1994 by Chapman & Hall, London. ISBN 0 412408201.
Farming systems 665
3. The source of water
4. The type of crops produced
5. Levels of production technology
6. Intensity of commercial production.
16.1.1 Land management systems

Five systems are generalJy recognized:
• Cropping plus fallowing

• Ley farming - alternate crop and livestock husbandry
• Field farming - permanent arable cultivation
• Perennial cropping - plantation style
• Agroforestry systems.
This first criterion in Ruthenberg's list is the most important basis for
classification of farming systems but the remaining elements also need to
be considered.
16.1.2 Intensity of cropping

This element is very closely related to section 16.1.1. Intensity of culti-
vation can conveniently be expressed in terms of an index, R, calculated in
terms of three variables:
R=~X100
d
where y = number of years under cultivation, c = number of crops per

annum, and d = duration of land-use cycle (crops + fallow).
Ruthenberg makes extensive use of this index and uses it to character-
ize farming systems:
• Shifting cultivation: R <30 - fallows are long.

• Semi-permanent cultivation: R = 30-70 - fallows relatively short.
• Permanent cultivation: R >70 - fallows very short or occasionaL
16.1.3 Water supply

The water needs for cropping can be supplied by:
• natural rainfall; or
• irrigation; or
• a combination of both.
16.1.4 Type of crops produced
Crops are produced broadly for four purposes:
• Food
• Beverages
• Industrial use
• Medicinal.
Individual crops may fall into more than one of these categories.
16.1.5 Level of production technology

Technological levels of farming systems can be considered in terms of
increasing levels of technological input and sophistication.
• Planting or digging-stick systems - the basic level and most primitive.

• Hoe or spade farming - representing a simple technologi~al advance
with human labour being the major source of energy input.
• Draught implements with animal power - the development of more
sophisticated equipment and technology with the supplementation of
human with animal power in energy input.
• Draught implements with mechanical power - producing a further
reduction in human energy input and replacement of animal energy by
that derived from fossil fuels.
16.1.6 Degree of commercialization

This is a pragmatic breakdown of what is clearly a continuum, compared
with 16.1.2 (intensity of cropping):
• Subsistence farming: sales <25% production.

• Partially commercialized farming: sales 25-50% of production.
• Semi-commercial farming: sales 50-75% of production.
• Commercial farming: sales >75% of production.
16.2 CULTIVATION SYSTEMS
The criteria in sections 16.1.2-6 superimposed on 16.1.1 give an adequate

basis for describing and interrelating different farming systems and the role
that the groundnut actually performs in them. For purposes of discussion,
systems of cultivation can be considered under seven different headings,
the first five of which will be discussed in detail:
Cultivation systems 667
1. Shifting cultivation
2. Semi-permanent cultivation
3. Ley farming
4. Permanent cultivation on natural rainfall
5. Permanent cultivation dependent in part or whole on irrigation
6. Plantations
7. Agroforestry systems.
The complexity of these systems is considerable and there are environ-
mental and other constraints which determine whether or not they are
practised in a given area and the precise form they take.
16.2.1 Shifting cultivation

This system is profoundly misunderstood in the western world, where land
is generally regarded as the most precious environmental resource and
anything but its most intensive use is considered profligate. This point of
view, though entirely understandable in those brought up in the developed
world, is certainly not tenable in the developing world and it would be
misguided in the extreme to maintain such a standpoint in these areas.
At first sight, shifting cultivation is aesthetically and intellectually un-
appealing, but on further examination it is revealed as astonishingly effec-
tive. It has been dismissed in a somewhat pejorative fashion as a
'technology of expediency'. There is undoubtedly an element of truth in
such a label but what practical alternative is there in vast areas of Africa,
Asia and South America? It is also a viable agricultural system over a wide
climatic range, from tropical rain forest to semi-arid environments. In
these areas the system works well provided that the alternating cropping
and fallowing phases are in balance. The essential requirement is that the
fallow period is of sufficient duration to restore soils and vegetation to the
state in which a further cycle of cropping can be supported. Depending on
soil type and present and past climatic conditions, the fallow period might
be quite short or extremely protracted. The practice of shifting cultivation
in the rainforest regions or of 'chitemene' in the woodland areas of Central
Africa may require something approaching the re-establishment of climax
vegetation before the cultivation cycle can be repeated (Trapnell, 1953;
Trapnell and Clothier, 1957; Allan, 1967). A great depth of ecological
perception is often apparent in the operation of these systems - in knowing
when the succession has reached the stage at which cropping can take
place.
The system works very well when populations are relatively low and
within the capacity of the environment to support. When population
pressures mount, there is an irresistible tendency to attempt cropping
before the soil has recovered fully from the previous cycle. Recovery of the
soil has physical and chemical components. In the fallowing phase, the
growth of grasses - especially their fibrous root systems - rebuilds the
physical structure of the soil, improving aeration and water acceptance, so
that field capacity is enhanced while allowing drainage of excess. Nutrients
released by weathering of soil minerals, and through activities of mycorr-
hiza and nitrifying bacteria, are recycled in situ and accumulate until an
equilibrium is established, at which point the cropping sequence can be
taken.
Fallowing may also be resorted to when problems of weed infestation
occur together with pest and disease build-up. This can happen on in-
herently fertile soils of good physical structure and high intrinsic nutrient
levels. There may come a time when the energy expenditure in maintaining
clean cultivation on an established plot is greater than that involved in
clearing a new one. It is primarily a question of labour input in relation to
the crop produced. In the case of the most primitive farming systems (using
a digging or planting stick, for example), the really significant inputs are
labour and seed or other propagating material which represents the capital
element. In the technologically more advanced systems of hoe and spade
culture, the capital element is somewhat higher. The substitution of animal
or mechanical power for human labour greatly increases the level of capital
commitment and associated risk.
Primitive agricultural systems (in the technological sense) using shifting
cultivation are in fact very much more efficient in agronomic terms and in
the use of environmental resources than appears at first sight to those
conditioned to different and more intensive agricultural practices. They are
also efficient in economic terms - that is, in the return on the labour
investment. Furthermore, the lifestyle which can be followed offers con-
siderable leisure time: the working day may be no longer than five hours on
average and peak demands for labour may be relatively few, apart from
sowing and harvesting. An important point is that more intensive systems
may demand a greater labour input for no greater return. The most
important point to appreciate in attempting to increase the intensity of land
exploitation is that increased levels of inputs are required, initially in the
form of plant nutrients, in order to increase levels of production. These
inputs must result in a clear surplus over subsistence needs which must be
adequate to cover their cost when marketed. Encouragement of increased
input levels without a reasonably assured saleable surplus can lead to
extremely serious problems of indebtedness, since farmers are essentially
living off capital which they do not actually possess. If they have title to the
land they cultivate, then they may well lose it in order to pay their debts.
16.2.2 Semi-permanent cultivation

This system, in which intensity of land use ranges from R values of 30 to 70,
is found in the seasonally dry to semi-arid tropics rather than in the humid
areas. Holdings tend to be more clearly defined; land occupancy by the
farmer and family may be of an indefinite term. The actual cropping
patterns and practices may be similar to those of shifting cultivation
practised in the same geographic area.
Semi-permanent cultivation is efficient and effective when a relatively
short fallow period is sufficient to restore fertility and reduce weed prob-
lems. However, where such a system ofland-use is enforced by population
pressures, the fertility and general condition of the land may become
impoverished, with continuing depression of yields and increased erosion
hazards. In such situations, research and extension operations should focus
on income generation once subsistence needs have been met so that
necessary inputs to maintain fertility and hence productivity can be
afforded. In these conditions groundnut could be highly desirable as a
valuable subsistence-cum-cash crop. The labour demands tend to be higher
under this system; the weed problems are usually more severe and little
in the way of land preparation may be possible during the dry season
(as would be the case with shifting cultivation). Peaks of labour demand
tend to be higher.
16.2.3 Ley-farming systems

After a period of popularity in the 1950s and of eclipse in the 1960s, ley-
farming or alternate husbandry systems are poised to stage something of a
comeback. Part of the reason is the increasing popularity of the ideas, at
any rate, of organic farming. Ley-farming systems are not of necessity
wholly organic - there is no reason why artificial fertilizers should not be
applied or agrichemicals used. However, there is invariably a substantial
cycling of organic matter within the system. Nutrients removed from the
farm by sale of crops and livestock must be replaced in order that the
system is sustainable at a reasonable level of productivity.
Ley-farming or alternate husbandry can be considered as being derived
from the semi-permanent system by the exploitation of the fallow for
grazing. In the primitive or unregulated form of ley-farming, the grazing of
livestock (where kept) is almost inevitable if grasses predominate in the
fallow vegetation when cultivation is abandoned. This system can be
developed or regulated to involve the deliberate sowing of grasses or a
grass-legume mixture in the ley. In the unregulated system, there need be
little capital investment over and above that required for semi-permanent
cultivation. The regulated system, however, can become fairly capital
intensive. In order to control grazing effectively, some kind of fencing is
required. At the time when the ley's useful life is at an end and it has to be
ploughed, investment in an effectively designed plough for the purpose
might well be necessary. Draught power could be provided by beasts
produced in the system.
The cropping system in the arable phase could take into account not only
the subsistence needs of the farmer (and the local community) but also
suitable crops for marketing in the immediate hinterland and those which
can be produced for export markets. On the livestock side, apart from the
draught animals already mentioned, the produce of a ley-farming system
would include meat or dairy products or both. In the tropics, fresh milk
production would imply a need for a site close to an urban population
which would provide a market. Meat production would not be subject to
this particular constraint if live animals could be marketed; otherwise there
would be a need for an abattoir, cold storage and refrigerated transport in
the vicinity.
Groundnuts could playa useful role in such a system. After harvest, the
haulm could be used as a very acceptable livestock feed and, if the seed
produced was used for oil extraction, the cake could be returned to feed
cattle and its nutrient content restored in part to the soil.
In theory this system seems to be well nigh perfect - a model of
sustainability, in fact. In the cropping phase soil nutrients are depleted,
physical condition of the soil deteriorates and weed populations may well
build up. Under the ley phase, arable weeds will be suppressed, soil
structure improved by grass root growth and the nutrient status enhanced
by the activity of the root-nodule bacteria of pasture legumes as well as the
dung of livestock, particularly if the animals are fed on imported concen-
trates. There is intrinsically a very good cycling of nutrients and organic
matter.
Given the right economic conditions, this system can be very attractive -
particularly in temperate climates, for which excellent grass-clover
mixtures are available. The system has been viable in New Zealand and
contributes largely to the country's exports of meat and dairy products.
There are problems, however, in non-temperate zones. At lower elev-
ations in the tropics, it is difficult to establish and maintain stable grass-
legume mixtures in pastures. A further problem is that the nutritive value
of tropical grass species is appreciably lower than that of the temperate rye
grasses. Tropical grasses tend to be lower in dry matter when they are
young and succulent and also tend to become fibrous more quickly with
age.
Although in ley-farming the land is being used productively all the time,
when pressures mount for maximization of production or income a greater
return can be obtained from continuous arable cultivation. For a ley-
farming system to be successful, population pressures in actual or potential
ley-farming areas have to be such as to encourage a higher level of
productivity but not so high that only continuous cultivation will meet
subsistence and other needs. The system also demands a high level of
farming skills and versatility: the farmer must be equally good with animals
as with crops and must achieve self-sufficiency of forage production. In
addition there must be no serious endemic disease problems and good
veterinary services should be available when needed.
These constraints are very severe in many if not most parts of the tropics
and preclude extensive development of the ley-farming system. Its intrinsic
intellectual and aesthetic appeal is considerable and it is well worth bearing
in mind under appropriate economic conditions.
16.2.4 Permanent cultivation on natural rainfall

This system can arise when arable cultivation is extended at the expense of
fallow or grass ley but it does include systems in which short-term fallows
or leys occur from time to time. A distinguishing feature of permanent
agriculture is the dedication of land on a permanent basis for arable or
grazing use. The arable land tends to be that which is most productive and
easiest to manage, while the grazing lands are most often those with
difficult topography, poor drainage or rocky outcrops which make them
unsuitable for arable crop production. In the seasonally dry tropics, annual
(sometimes biennial) crops are taken under arable systems; in the wet
tropics, plantation tree crops are customary.
The most satisfactory systems of permanent cropping develop in areas of
high intrinsic soil fertility, where this does not decline rapidly or drastically
under cultivation. Such areas are those with young soils of volcanic origin
or those developed in Rift Valley areas following the causative earth
movements. They are characterized by seasonal rainfall or well-distributed
rainfall in which leaching is not severe. This system is most readily main-
tained where fertilizer use is established on an intensive basis - a practice
which maintains permanent cultivation in advanced agricultural econo-
mies, where intensive fertilizer use is paid for as a charge on the crop.
Without this high level of input, it is questionable whether intrinsic fertility
would sustain cropping at economic levels for any great length of time. In
western Europe over the past two centuries, fertility has been imported not
only in the form of N P K fertilizers but also in livestock feeds, often
imported from the developing world - such as groundnut cake produced
from oil extraction.
Permanent agriculture in the tropics is quite commonly poles apart from
that found in the developed agricultural economies. In the Indian subconti-
nent, for example, population pressures may maintain land in permanent
cultivation which would be better managed under a system with fallows.
Inputs of fertilizer and manure are commonly inadequate and often negli-
gible. Cow dung in India is too valuable as a fuel to be used as manure,
while in Africa there is frequently no established tradition of using cattle
manure. Farmers, particularly in tsetse-infested areas, have no experience
of cattle management; on the other hand, some herdsmen have never
produced crops. Agricultural development has often brought tsetse under
control, enabling farmers to keep cattle, and settlement schemes have
introduced pastoralists to agriculture. Both groups have to acquire the
necessary skill in the handling of manure and this requires investment in
carts if manure has to be transported for any substantial distance.
A general characteristic of permanent cultivation in the tropics is that
demand for labour per unit area is high and that output may be relatively
low in impoverished or degenerate systems. Ruthenberg (1976) described a
system of agriculture developed by the Wakara people on an island in Lake
Victoria. As population grew, they were unable to expand the area under
cultivation and had perforce to intensify their agriculture. They developed
a system in which fertility resources of all kinds, notably animal and green
manures, are used to the full to maintain their land at an adequate level of
productivity. The labour demand for this system is very high and those who
have emigrated from the island and who continue to farm readily abandon
their remarkable system of cultivation: they are very happy to adopt semi-
permanent or shifting agricultural systems, which are much less laborious.
The farming of the Wakara is particularly interesting because it shows the
development of intensive land use comparable to that of many peoples of
the Far East.
The role of the groundnut in such systems of permanent cultivation is
basically similar to the role that it performs in fallowing or ley-farming
systems. There is, however, the constraint that the high labour demand of
this crop may sharply limit the extent to which it can be grown. Since it is a
highly regarded food crop, it is likely that it would be grown anyway in
suitable climatic areas on appropriate soil, even if only to a small extent; on
the other hand, if a good local market exists it could be useful as a cash-
earner, again even if only to a limited extent. In such areas it would
obviously not be easy to expand production without some additional
technological input.
16.2.5 Permanent agriculture under irrigation

In many ways this system represents an ideal. By implication it is practised
in arid areas where leaching is minimal and fertility is high and the crop's
needs for water can be met exactly, assuming that water supplies are
constantly available. But for irrigation, some highly productive agricultural
areas such as Egypt and California would be desert. In these areas tem-
peratures are not limiting, although the diurnal range may be quite wide.
The important considerations are availability of water when needed and its
quality. Without appropriate care and attention, solute accumulation can
occur to the detriment of future production. The maintenance of irrigation
agriculture in the Nile Valley shows that such systems can be sustainable
indefinitely. Problems can arise when attempts are made to expand culti-
vation with a limited water supply, which can bring disaster in their wake,
or where there are drainage problems due to local topography.
Groundnuts have been produced successfully in the irrigation agriculture
of Egypt for many years. The crop lends itself well to such a system. It has
clearly defined critical periods when adequate water supplies are necessary
for peak yield, the most important (after germination and establishment of
the crop) being from flowering until pod-filling is complete. There are
other situations where irrigation can supplement natural rainfall to ensure
an adequate length of growing season. In semi-arid tropical areas, irriga-
tion may be used to prolong the growing season by establishing crops prior
to the onset of the rainy season, a practice which also gives the crop an
extended period of freedom from the normal pests and diseases of the
growing season. On both counts, yields can be increased without the cost
of agrichemicals and their undesirable environmental effects. Irrigation
can also be used at the end of the season to facilitate crop maturation when
soil water has become limiting. ,
In certain areas the groundnut crop can be linked with production of
rice, particularly in the humid tropics. In many parts of the wet tropics,
rainfall can be expected at any time of the year but distribution is not
uniform and it may be possible to combine production of paddy rice at the
wetter times of the year with groundnut production in the 'dry' season.
During this period less difficulty will be experienced with the drying and
curing of the crop than in periods of the heaviest rainfall.
Groundnuts can be grown on quite heavy soils, provided that they are
ridged and other suitable practices are adopted. They have been grown on
the Gezira scheme in the Sudan on heavy cotton soils. In fact more serious
problems have arisen in Africa from attempts to produce the crop on light
soils (the East African Groundnut Scheme) than on heavy soils.
16.2.6 Mixed cropping versus sole cropping

In traditional systems, groundnut has often been produced in some kind of
a mixture with other crops. Neither the components of the mixture nor
their proportions are fixed. Such systems have considerable advantages,
the chief of which is flexibility. In a subsistence economy, the balance
between the components can be determined on the basis of need. The
staple crop will naturally occupy the greater proportion of the land and the
amounts grown of subordinate crops can be varied according to need. The
mixture tends to be sown in a succession. In the mound culture systems of
Central Africa (Trapnell 1953), the mounds are often set up by burying
materials from land clearance or previous crop residues in the late dry
season or early rains, followed by sowing maize and sorghum centrally on
the mound and then a succession of other crops such as groundnuts,
cowpeas, cucurbits, sweet potatoes and Phaseolus beans. At the peak of
the season the ground cover can be excellent and erosion minimized.
This simple succession can lead to sowing of crops in a series of more or
less concentric circles, which is satisfactory in a hand tillage system. It can
be adapted to animal-powered tillage by producing a linear version. The
staple crop can be sown in widely spaced lines and, following each culti-
vation, additional components can be added to the system in amounts
674 The ground nut in farming systems and the rural economy
determined by need. This 'relay cropping' (as it is commonly called) can be
related to traditional farming practices adapted and modified to suit more
advanced technology.
There are additional advantages of traditional and modern systems of
mixed cropping in that they tend to reduce the incidence and severity of
pest and disease attack. In a pure stand the pest needs to expend little
energy in finding a host plant; similarly with plant pathogens the dispersed
inoculum has nothing to alight on other than host plants. In mixed crops
the effectiveness of pests and disease organisms in finding suitable host
plants is substantially reduced. Development of infestations and infections
is slower and epidemics are much less likely to occur. Also, the rooting
zones of the component species in the mixture are unlikely to coincide, so
that more effective exploitation of the whole soil profile is possible.
Appropriate plant spacing could minimize competition between the root
systems and allow the effective interception of light by the canopies. Light
shading may even benefit ground nuts at pegging. These factors could have
a cumulative effect and enhance overall yield relative to what the separate
components might have achieved in pure stands from an equal area of
land.
In traditional systems where mixed cropping is standard practice, some
crops - such as finger millet (Eleusine coracana) - are invariably produced
in pure stands. This is usually related to ease of harvesting of the small
grains. When cash crops are introduced to such agricultural economies,
they are frequently grown as pure stands even when they are part of a
traditional system (as groundnuts are). Several considerations lead to the
adoption of this practice in such circumstances. In the case of tobacco and
cotton, ease of harvesting is an important consideration; in addition, in
cotton, economic use of pesticides would be difficult in mixed crops even
though the pests themselves might be less of a problem in such
circumstances.
The change from mixed to sole cropping has been hastened by use of
animal and mechanized power in crop production. It has without doubt
increased the biotic pressure of pests and pathogens on the crops, so much
so that interest has revived in mixed cropping systems such as relay
planting and agroforestry. They provide alternatives which seem to be
more environmentally friendly as well as meeting the subsistence needs of
the farmer more effectively. There is no reason why cash crops could not
be produced by relay planting and from agroforestry systems where, for
example, no restrictive considerations with respect to harvesting were
involved.
The theoretical basis for agroforestry systems is similar to that of relay
cropping. The principal difference is in the time scale: relay cropping
typically involves cropping one season at a time while in agroforestry a
long-term perennial element is combined with production of annual crops.
Agroforestry could also be regarded as an indefinite extension of the
Adaptive research 675
system used in establishing a plantation of tree crops when annual or other
short-term crops are taken before the plantation comes into production.
16.3 ADAPTIVE RESEARCH
The subject of mixed cropping, intercropping, multiple cropping or poly-

culture has received some attention in the past two decades in the devel-
oped world. The American Society of Agronomy published its symposium
on multiple cropping in 1976. Among nume,rous other publications in the
area, one can mention Vandermeer (1989), who develops a detailed theor-
etical consideration of the subject. This is also considered in the broader
ecological context by authors such as Tivey (1990) and Loomis and Connor
(1992). The subject is one of some complexity, as Vandermeer's (1989)
work shows. In practice, it is necessary to develop an understanding of the
needs of individual crop species for optimal production before fully effec-
tive manipulation of multiple cropping can be achieved.
In the context of improving production of the crop in the developing
world, or in its introduction to new areas, it makes good sense to use
existing knowledge of the crop's production to the full. and to draw upon
experience with the crop in technologically advanced areas. However, it is
foolhardy to proceed without preliminary exploratory trials - in other
words, to undertake what is now called adaptive research. There is a real
necessity to carry out locally-based programmes to determine how best to
apply existing knowledge to other environments and how to fine-tune and
adapt them to different locations.
The East African Groundnut Scheme provides a perfect illustration of
what can go wrong when no prior adaptive research is carried out. From
experience of the crop in North America and South Africa, it was believed
that certain short cuts could be taken; it was assumed that lighter soils
would suit the crop and that the short season cultivar Natal Common was a
sensible choice. As it transpired, the light soils were a very different
proposition from those of North America - instead of being loose and
friable when dry, they set almost like concrete. It was most difficult to
harvest the crop and the wear and tear on harvesting machinery was totally
unacceptable. On the face of it, Natal Common was a reasonable choice-
as a spanish variety it is quite early and capable of maturing in a relatively
short growing season - but no pilot trials were conducted to determine
whether in fact any groundnut genotype, no matter how early maturing,
could succeed in the conditions at Kongwa.
16.3.1 Adaptive research in Zambia

In contrast to the East Africa scheme, an adaptive research programme did
evolve in Zambia in the period 1954-61. It began as a breeding and
selection programme. After the third season, when promising germplasm
had been identified, it was apparent that available local knowledge was not
such that the extension service could advise as to how to make the best use
of any new material. The decision was made to undertake what would now
be called a programme of adaptive research, the results of which are given
by Smartt (1967). The outcome was a sound crop production package for
the grower.
Three or four decades later, that approach is still relevant to any area
where new productive varieties are being sought and where it is necessary
to determine how to make the best use of any improved varieties that have
been produced or identified. The following is an extended summary of the
Zambia programme.
The groundnut crop in Zambia had been produced with varying success
in different parts of the country. In the Eastern Province there was an
export of high quality confectionery kernels, mainly from the variety
Chalimbana (a Jumbo type). Production standards were high and yield
levels were quite satisfactory. It was not clear why yields were so much
lower outside the Eastern Province or why the crop was not a very
attractive proposition as a generator of income elsewhere.
(a) Variety selection

A series of variety trials, run for four seasons from 1954 to 1958, included
four locallandrace varieties (Ngoni, Nsenga, Chewa and Chalimbana), the
standard South African cultivar Natal Common and an introduction from
Tanganyika (Asiriya Mwitunde). In the first season, 1954-55, the trial was
sown at only one location - Mukulaikwa, the site of an abandoned
Overseas Food Corporation pilot groundnut scheme. Yields in this first
season were abysmal (Table 16.1); in the second season they were worse
and no further groundnut trials were carried. out on this site after 1956. In
the season 1955-56, experiments were carried out at two sites with the
same varieties, one in the Eastern Province the other in the Southern
Province. While the yields produced in the Eastern Province (Kalichero)
were marginally better than those of the Southern Province (Monze), they
were encouragingly close and gave an indication that acceptable, economic
yield levels were possible elsewhere than in the Eastern Province. In
1956-57 the trials were again repeated in the Southern and Eastern
Provinces and in this season the performance at Monze was actually
considerably better than that at Petauke (Eastern Province). In the final
season of these trials, the tables were turned and yields at Petauke were
highest while those at Monze were among the lowest. In this last season,
the trial was carried out at six locations with indications that at Kabompo,
for example, there might be useful production potential.
An introduction nursery had been established at the Central
Agricultural Research Station in 1954 and it became apparent in its early
TABLE 16.1 Mean yields (kernels lh/acre) of groundnut varieties, 1954-8
Season Station Province Ngoni Nsenga Chewa Chalim- Natal Asiriya SE LSD
bana Common Mwitunde P = 0.05
1954-5 Mukulaikwa Central 110 140 225 200 50 240 ±22 69
1955-6 Mukulaikwa Central 51 164 82 193 38 216 ±33 99
Kalichero Eastern 795 870 906 872 710 900 ±39 120
Monze Southern 813 783 776 872 588 680 ±38 116
1956-7 Monze Southern 1375 1188 1243 1254 957 1111 ±32 97
Petauke Eastern 363 594 583 594 286 • 539 ±24 75
1957-8 Petauke Eastern 1232 1243 1199 1122 1078 704 ±34 104
Katapola Eastern 693 836 792 748 429 616 ±68 205
Monze Southern 429 484 418 429 264 429 ±41 125
Misamfu Northern 660 693 649 726 528 660 ±69 207
Balovale N. Western 407 396 429 275 33 374 ±41 124
Kabompo N. Western 957 957 671 880 143 792 ±72 219
TABLE 16.2 Mean yields (kernels lh/acre) of groundnut varieties, Mount Makulu,
1956-9
Season Variety
mean yields
Variety Habit 1956-7 1957-8 1958-9 1956-9
Mani Pintar Bunch 2192 1011 1472 1558

S183 Runner 1841 771 995 1202
GB1 Bunch 1738 846 1095 1168
Dixie Runner Runner 1621 938 404 988
BS1 Runner 1603 696 741 1013
S731 Runner 1589 950 736 1092
Chitedze Mwitunde Bunch 1512 631 630 924
TMV3 Bunch 1509 743 609 954
Asiriya Mwitunde Bunch 1441 602 691 911
Seasonal mean yields 1672 799 819
Standard error of
variety means ±102 ±49 ±58
LSD P = 0.05 299 158 190
years that some introductions had a considerably greater yield potential

than either the locallandraces or introductions received from South Africa
or Tanganyika. In the season 1956-57, sufficient bulk of material had been
produced to undertake a new series of variety trials which were carried out
from 1956 to 1959. These had only one variety (Asiriya Mwitunde) in
common with the earlier series of trials. They showed that the introduced
Bolivian variety Mani Pintar (undoubtedly a corruption of mani pintado)
was the highest yielder in three very different and successive seasons. The
other conclusion which could be drawn from the yield data (Table 16.2)
was that, although many varieties performed well in the excellent 1956-57
season, there was little consistency of performance (apart from Mani
Pintar) over all three seasons. It was possible to identify at a relatively
early stage germplasm which showed promise and consistency in
performance.
A consideration of the results of the earlier trials (Table 16.1) revealed a
lack of consistently good performers but Natal Common was consistently
poor. Of the six varieties tested, Chalimbana was generally preferred by
growers and COnsumers on the grounds of its excellent quality and because
in yielding ability it was marginally higher and more consistent than the
others. In the planting of these trials (at the provincial sites) it was
requested that they be sown at about the same time as the local crop. From
nursery experience at Mount Makulu and anecdotal evidence from the
Eastern Province, it became apparent that the crop benefitted greatly from
early sowing. When promising lines had been identified the need was felt to
carry out agronomic investigations to determine how best to realize the
potential for high consistent yield shown by Mani Pintar.
The variety Mani Pintar has an interesting and chequered history. It was
collected in 1947-8 (Hartley, 1949) at a local market in La Paz, Bolivia. It
was sent to the United States and to Australia, where it was grown
experimentally in Queensland. A small sample was received in Zambia in
1955 and grown in nursery plots. Single plant selections were made and the
remaining bulk was used for sowing in variety trials from 1956 onwards.
Small samples were sent elsewhere in Africa and promising results were
reported from Ghana initially (McEwen, 1961), Zimbabwe (Smartt, 1978;
Hildebrand and Smartt, 1980), East Africa and as far north as the Sudan.
Extensive trial sowings were made throughout Zambia and promising
results and commercial bulking commenced in 1960.
The original material of Mani Pintar showed very striking variegation in
pigmentation of the testa. This was thought to be a possible disadvantage
in terms of marketing. Single plant selections were made of plants which
produced seeds with self-coloured testa, both red and brown. The red-
seeded selection bred reasonably true and was named Makulu Red; the
brown-seeded material segregated and was clearly hybrid in nature. The
behaviour of both Makulu Red and Mani Pintar when produced in bulk
was interesting in that it was usually possible to select some self-coloured
seed in a variegated batch and some variegated seed from a self-coloured
batch. Since the two forms were otherwise identical, it was suggested that a
position effect might be responsible (Srtlartt, 1960). Hammons (1973)
showed that variegation behaves as a single locus effect but an alternative
explanation is that the difference is due to a jumping gene or transposon.
This possibility might be worthy of further study and it could, if establis-
hed,have interesting implications.
Further selection in Zimbabwe (Hildebrand and Smartt, 1980) has
shown that stable pink and brown testa forms can be selected from Mani
Pintar and Makulu Red stock. The potential value of Bolivian germplasm
in Central Africa (and elsewhere in the continent) has been suggested. This
value derives from the fact that it evolved in a montane environment,
probably at about 1500 m a.s.l., where day temperatures may be high with
cool conditions at night. These conditions are in marked contrast to those
which occur in many of the major growing areas closer to sea level, where
temperature range is much less, and it seems likely that rather different
physiologies have become established. Conditions in plateau Africa prob-
ably have a great deal in common with those in which Bolivian groundnuts
have evolved. What is more certain is that they show remarkable pre-
adaptation to conditions over much of Africa.
From a commercial standpoint, Mani Pintar (and Makulu Red) have
certain good features and some shortcomings. The size and shape is
relatively uniform but somewhat shorter in relation to diameter than in
typical virginia varieties. The oil content is about 50% of dry matter; the
kernel flavour is somewhat bland and thus it is better used where a strong
flavour is not required, rather than for the preparation of salted or roasted
peanuts. However, their potential value in breeding programmes is high.
They have a further advantage in that their maturation period is intermedi-
ate between that of a typical virginia and a typical spanish/valencia type.
This has proved to be particularly valuable in Central Africa, where
virginia types require a somewhat longer maturation period than is avail-
able in many seasons, while the fastigiate valencia/spanish cultivars mature
before the rains finish and losses may occur from sprouting. In Zambia the
maturation period was found to be about 10 days less than the typical
virginia and 20 days longer than spanish/valencia. A crossing programme
between Mani Pintar and Chalimbana, although it was never launched,
could have had exceptionally interesting results if it had proved possible to
combine the confectionery quality of Chalimbana and the yielding capacity
of the Bolivian germplasm.
(b) Agronomic practices

The identification of high-yielding and well-adapted genotypes is essential
but it is only the first part of the task of the crop improver. A good
genotype grown in poor conditions will not perform well, neither will an
unproductive genotype grown in the best conditions that can be provided.
The breeder initiating a selectinn programme in a new area will naturally
look to locallandrace material as well as introductions to provide the pool
of variation from which to select. Locallandraces are usually considered to
be adapted to local conditions and this is frequently so in ways that are not
appreciated. One would expect them to be attuned to local climatic
conditions and those of the local environment generally. This might well
include low levels of fertility: they may be able to produce a modest crop
annually with negligible inputs of nutrient but they are commonly unable
to respond effectively to substantially improved conditions. As local lan-
draces they have never previously experienced such conditions and have
been subjected to no previous selection for such responses, so that it should
be no surprise if they do not respond markedly to improved cultural
conditions.
In the course of nursery operations and carrying out the initial series of
variety trials, the following were identified as the most important areas for
agronomic investigation:
1. Time of sowing
2. Time of lifting
3. Plant population and spacing
4. Cultivation and weed control
5. Fertilizer use.
In addition, the effects of disease and (to a much lesser extent) pests were
investigated.
1. Time of sowing
This was identified in the very first season as an important factor influenc-
ing yield. The initial nursery collection was sown in duplicate, the first
replicate as soon as was practicable after the first rains in mid-late
November and the second replicate approximately 4 weeks later in mid
December. At harvest it was quite obvious that the early sowing had out-
performed the later by a considerable m~rgin and it was thought that an
attempt to quantify the effects of delayed sowing would be a useful
exercise. It soon became apparent that general practice among local tradi-
tional farmers was to sow their most important crops as early as possible.
In the Eastern Province, where groundnuts were an important source of
income, this category included groundnuts which could well be sown
before 1 December. In other areas,where they were solely a subsistence
crop, planting tended to be later after the sowing of staple crops - maize,
sorghum or finger millet - and this could be as late as mid December.
Sometimes, if onset of the rains was delayed, the earliest sowing might well
be possible only as late as mid December.
Results from trials conducted in two seasons are presented in Tables 16.3
and 16.4. These show consistent trends, with delayed sowing producing
substantial yield reductions. In the case of the Mount Makulu trials, where
a kernel quality test was also carried out, reduction in quality follows a very
similar trend to yield. A general recommendation can therefore be given,
on the basis of these (and other similar) results, that advantage be taken of
the earliest possible opportunity for sowing. Sometimes a problem arises in
connection with the earliest sowings when moisture distribution is not
uniform in the field and uneven germination results, but the consequences
of delayed sowing are likely to be even more detrimental.
Another important effect of early sowing is to reduce the impact of some
major pests and diseases on yield. This will be considered in more detail
later in section 16.3.1(c).
2. Time of lifting
The question of harvesting a geocarpic crop with a fundamentally indeter-
minate flowering sequence is a vexed one. It is a matter of determining the
optimum time when the highest proportion of pods are mature. Premature
harvesting will obviously increase the proportion of immature fruits har-
vested as well as giving a yield reduction, while delayed harvesting pro-
duces loss of early-set pods by breakage or disintegration of pegs, severing
the connection between plant and pods. Although all groundnuts are
botanically indeterminate in flowering and fruiting, in practice the fasti-
giate varieties and the spreading bunch forms tend to have a defined period
of effective flowering and fruiting. Only those pegs fairly close to the
TABLE 16.3 Effect of sowing times on kernel yield and quality of three groundnut varieties at Mount Makulu, 1959-60
Variety OBI Makulu Red Local spreading 2

Sowing date Mean yields % retained by Mean yields % retained by Mean yields % retained by
kernels 19/64" x 314" kernels 19/64" x 314" kernels 19/64" x 314"
lblacre screen lblacre screen lblacre screen
SE ±52 ±46 ±52
21 Nov. 690 17.00 1115 63.27 840 78.34
10 Dec. 761 26.44 828 75.90 707 80.02
23 Dec. 394 20.44 411 85.19 340 70.62
8 Jan. 109 17.74 154 67.07 88 56.89
21 Jan. 21 8.33 28 30.48 25 14.28
LSD P = 0.05 158 140 158

TABLE 16.4 Effect of sowing time on kernel yield at several stations in Zambia (mean yields, kernels lb/acre)
Season: 1959-60 1960-1

Station: Petauke Balovale Monze 1 Monze 2 Monze 3 Monze 4
Variety: Mani Pintar Local Runner
Makulu Red
Time of mid- mid- mid- mid- mid- mid-
1st sowing: December November November November November November
SE ±49 ±39 ±82 ±73 ±117 ±73
1st sowing 2513 729 1298 1528 2279 1611
2nd sowing 1531 489 1133 1288 1864 1324
3rd sowing 104 358 524 705 1467 827
4th sowing 82 228 559 541 837 661
5th sowing 17 120 245 416 562 358
LSD P = 0.05 146 110 251 228 360 230

taproot borne on the lower nodes of the stem actually mature fruit. Pegs
produced higher up the stem fail to make soil contact and develop into
mature fruit. Maturity of individual pods is indicated by a darkening of the
tissues on the inner pod wall. Since pods at varying stages of maturity are
to be found on a single plant, it is not always easy to gauge maturity of a
whole field crop from a few randomly selected plants.
It is important to know not only the optimal time for lifting but also the
latitude that exists and the effects of premature or delayed harvesting on
yield and quality. The question of quality these days embraces not only
that of the kernel but also levels of infection by Aspergillus flavus and A.
parasiticus and the resulting aflatoxin contamination. The most effective
way of elucidating the effects of time of lifting on yield and quality is to
carry out a quality test in addition to yield measurements soon after
harvest. This involves screening the crop and recording the proportion of
large kernels produced, a simple and effective quality assessment.
The results of a series of trials are presented in Tables 16.5, 16.6 and
16.7, which clearly show that the optimal lifting time for Mani Pintar/
Makulu Red is 150 days. There is a window between 150 and 160 days,
after which serious yield loss can occur. For earlier maturing cultivars
(Spanish GBI) the optimum appears to be 130 days; for the later maturing,
spreading varieties it is about 160 days. While these indications are reliable
on the whole, they can be affected by seasonal conditions. In particular,
dry conditions post-sowing could cause delayed emergence and if this is
longer than a few days due allowance has to be made in deciding when to
lift.
3. Plant populations and spacing

The initial study of the effect of plant populations included a number of
ridging treatments. These were recommended as good soil conservation
practices and the opportunity was taken to assess possible effects on yield
in a single season. The results presented in Table 16.8 show no significant
effects of either cultural practices or plant populations. The latter were
changed by varying density within rows; the rows themselves were on
average 36 inches (91 cm) apart.
In order to obtain a satisfactory understanding of the question, it was
clear that spacing between and within rows would have to be varied,
subject only to the constraint of what could be accommodated in a standard
size plot (25 ft x 10 ft). Experimental results are presented in Tables 16.9
and 16.10). The optimal plant population under the rain-fed conditions, in
which these experiments were conducted was the intended density of
43 560 plants per acre as far as both yield and quality were concerned. It
was interesting to note that yield fell less sharply than quality at higher than
optimal plant populations. The higher proportion of shrivelled kernels
suggested that, at the higher population densities, more pods were set but
that water availability was inadequate to take the kernels through to
TABLE 16.5 Effect of lifting on kernel yield and quality of three groundnut varieties at Mount Makulu, 1959-60
Variety: (a) GBI (b) Makulu Red (c) Local Spreading 2

Time to lifting (days) Mean yields % retained by Mean yields % retained by Mean yields % retained by
for each variety kernels 19/64" x 3/4" kernels 19/64" x 3/4" kernels 19/64" x 3/4"
(a) (b) (c) lb/acre screen lb/acre screen lb/acre screen
(SE ±27) (SE ±44) (SE ±79)
100 120 130 287 3.66 506 12.80 732 49.28
110 130 140 424 6.61 674 43.90 957 71.11
120 140 150 436 12.40 1183 69.18 817 79.44
130 150 160 495 19.43 1211 80.34 1120 84.08
140 160 170 518 15.88 1229 87.60 550 82.80
LSD P = 0.05 83 135 242

TABLE 16.6 Effect of time of lifting on kernel yield and quality of two groundnut
varieties at Mount Makulu, 1960-61
Variety: (a) GB1 (b) Makulu Red
Time to lifting Mean yields % retained by Mean yields % retained by

(days) for each kernels 19/64" x 314" kernels 19/64" x 314"
variety Ib/acre screen Ib/acre screen
(a) (b) (SE ±86) (SE ±74)
100 130 318 4.56 1088 59.28

120 150 619 24.88 1322 83.88
130 160 700 18.25 1177 80.13
140 170 503 24.16 837 80.65
150 180 348 33.05 721 82.06
LSD P = 0.05 264 226
TABLE 16.7 Effect of time of lifting on kernel yield of

groundnuts at two stations in Zambia, 1959-61 (mean
yields, kernels lblacre)
Variety: Mani Pintar Makulu Red

Station: Petauke, 1959-60 Magoye, 1960-61
Days to lifting (SE ±99) (SE ± 127)
120 2046 1181

140 2151 1590
150 2152 2010
160 1888 1971
180 1877 1277
LSD P = 0.05 301 390
maturity. The inference could also be drawn that additional water supplied
by irrigation could have taken more of the kernels to full maturity and
improved both yield and quality from the higher populations.
4. Cultivations and weed control

The effects of the number of cultivations on yield were studied in season
1958--61. Initially up to five cultivations were compared with an unweeded
control at three sites (Table 16.11). The unweeded control plots gave very
low yields and at least two cultivations were necessary at some sites (and
TABLE 16.8 Effects of ridging methods and plant populations on the yield of
groundnuts, 1958-9 (kernels lb/acre)
Station: Mount Makulu Ngoni Farm Petauke

Province: Central Eastern Eastern
Variety: Mani Pintar Local Runner Local Runner
Ridging treatments (SE ±74)" (±54)C (±40)C Mean
Flat 1061 483 552 699

Flat and ridged 1089 436 534 686
Narrow ridge 1067 536 499 701
Broad ridge 1094 576 525 732
Populations thousands/acre
(SE ±64)h (±46)d (±35)f
20 955 461 491 636
40 1103 505 515 708
80 1175 560 576 770
Mean 1078 509 527
Least significant differences P = 0.05

a b c d e f
230 199 169 149 107 126
TABLE 16.9 Effect of plant spacing on yield and quality of two varieties of
groundnuts at Mount Makulu, 1959-60
Variety: GB1 Makulu Red
Intended Plant Mean yields % Mean yields %

population spacing kernels retained by kernels retained by
plants/acre Ib/acre 19/64" x 3/4" Ib/acre 19/64" x 3/4"
(SE ±88) screen (SE ±117) screen
14520 36" x 12" 624 27.49 623 93.98

29040 36" x 6" 781 28.09 963 86.09
58080 36" x 3" 1030 20.68 1256 84.19
29040 18" x 12" 1116 26.86 1139 86.66
58080 18" x 6" 1329 17.64 1065 76.63
116160 18" x 3" 1333 8.95 1270 68.99
43560 12" x 12" 1443 21.56 1364 86.56
87120 12" x 6" 1290 6.81 1132 69.63
174240 12" x 3" 1268 6.64 1045 52.95
LSDP = 0.05 262 351
688 The ground nut in farming systems and the rural economy
TABLE 16.10 Effect of plant spacing on kernel yield of groundnuts at Monze,
1960-1 (mean yields, (kernels lblacre), variety Makulu Red
Station: Monze 2 Monze 3 Monze 4

Intended Plant spacing (SE ±42) (SE ±163) (SE ±102)
population
plants/acre
21780 24/1 X 12/1 1341 2119 798

43560 24/1 X 6/1 1400 2226 1197
43560 12/1 X 12/1 1524 2562 1391
87120 12/1 x 6/1 1493 2579 1352
LSD P = 0.05 125 492 310
TABLE 16.11 Effect of number of cultivations on the yield of groundnuts, 1958-9

(kernels lb/acre)
Station: Mount Makulu Ngoni Farm Petauke

Province: Central Eastern Eastern
Variety: Makulu Red Local Runner Local Runner
Number of cultivations ±104 ±61 ±69
0 137 228 63
1 278 468 489
2 1053 762 440
3 1162 936 599
4 1489 980 651
5 1467 893 600
LSD P = 0.05 313 189 214
even more in some seasons) to approach maximum yield levels. There

were additional benefits for up to four cultivations, even though the
amount of weed growth removed was relatively small. Late in the season,
competition between weeds and crop for water may produce some loss of
yield and quality when water supply becomes limiting. When the trial was
repeated in the following season (1959-60) at Mount Makulu (Table 16.12)
there was a progressive increase in both yield and quality for every
cultivation given. In provincial trials (Table 16.13) the results of Petauke
and Balovale show an interesting contrast. Yield at Balovale is unrespon-
sive to cultivation, reflecting very low weed populations, while the pattern
of response at Petauke closely parallels that observed at Mount Makulu.
TABLE 16.12 Effect of cultivations on groundnut
yield and quality at Mount Makulu, 1959-60, variety
Makulu Red
Number of Mean yields % retained by

cultivations kernels 19/64" x 314"
lb/acre screen
(SE ±65)
o 481 56.72
1 620 58.91
2 821 65.03
3 914 68.59
4 1139 71.43
LSD P = 0.05 195
TABLE 16.13 Effect on cultivations on yield at two

stations in Zambia, 1959-60 (mean yields kernels
lb/acre)
Variety: Mani Pintar Local Runner

Station: Petauke Balovale
Number of (SE ±77) (SE ±59)

cultivations
0 828 827
1 1683 849
2 2090 914
3 2132 870·
4 2410 837
LSD P = 0.05 233 180
In the final season of experimentation at Mount Makulu a rather differ-

ent approach was adopted, although an uncultivated control was also used.
The crop was allowed to compete with weeds for varying periods and then
clean weeded thereafter (Table 16.14). The results show that three weeks
of weed competition between 6 and 27 January had little effect on yield and
none on quality. Longer delays produced very substantial yield losses but
very little reduction in quality. This result is particularly interesting be-
cause peak rainfall in Zambia usually occurs in January and there is a
temptation to cultivate even though the soil may be in an unsuitable
TABLE 16.14 Effect of withholding cultivations on
yield quality of groundnuts at Mount Makulu, 1960-1,
variety Makulu Red
Date of first Mean yields % retained by

cultivation kernels 19/64" x 314"
Iblacre screen
(SE ±61)
6 January 1011 79.35

27 January 933 80.47
10 February 440 78.15
10 March 172 71.19
Uncultivated 92 59.10
LSD P = 0.05 190
condition. This result indicates that weeding might possibly be deferred

without significant loss of yield until soils are less waterlogged and culti-
vation is likely to be more effective.
5. Fertilizer use
In what was current American practice during the 1950s and 1960s, the
only direct fertilizer applied to the groundnut was gypsum to ensure
supplies of calcium adequate for full normal pod development. While there
is no doubt that groundnuts are responsive to soil fertility as generally
perceived, responses to fertilizers in the season of application are inconsist-
ent and erratic. This conclusion is supported by trials carried out in Zambia
in 1958-59 (Table 16.15). It can be argued that the crop is in fact not
responding to the fertilizers applied, since these are in all probability not in
the same soil horizon as the feeding roots of the crop. This supports the
wisdom of the old American guideline to fertilize the rotation as a whole
rather than the groundnut break. Nutrients which have passed beyond the
reach of the feeding roots of maize, for example, may be readily accessible
to the groundnut root system. It would have been interesting to follow the
effects of deep fertilizer placement.
(c) Pest and disease control

For local farmers to use pesticides and fungicides was not a practical
proposition. However, the possibilities of cultural control were thought to
be worth exploring. The two major disease problems studied were rosette
virus and the cercospora leaf spots which, although produced by two
different pathogens (Cercospora arachidicola and Phaeoisariopsis perso-
nata), can conveniently be considered as a single disease.
The rosette virus is transmitted by the aphid (Aphis craccivora) (Chapter
TABLE 16.15 Responses of groundnuts to fertilizers, 1958-9 (kernels lblacre)
Station: Mount Magoye Masaiti Petauke Misamfu

Makulu
Province: Central Southern Western Eastern Northern
Variety: Mani Pintar Mwitunde Local Runner Local Runner Mwitunde
Effects ±37 ±117 ±139 ±20 ±26
N* 142 -104 135 -87 200

pt -133 27 85 -52 176
K++ -78 -12 -50 -17 54
Mean yield 1760 1351 602 862 804
LSD P = 0.05 114 360 424 61 80
* Response to 100 lb ammonium sulphate per acre

t Response to 200 lb single superphosphate per acre
++ Response to 100 lb muriate of potash per acre
10) and there are two avenues of attacking the problem: controlling the
insect vector by host-plant resistance and/or by resistance of the host plant
to the virus itself. At the time of this experimental work, host resistance to
the aphid had been found in East Africa in the variety Asiriya Mwitunde.
Field tests at Mukulaikwa in Zambia showed that, under field conditions in
which 100% infection was observed in Natal Common, only 2% occurred
in Asiriya Mwitunde. Clearly antixenosis was operating, the basis of which
has not been explored. Evans (1954) pointed out the difficulty of carrying
out experimental studies on rosette using conventional small plot tech-
niques. This difficulty was also experienced in Zambia but, very occasion-
ally, analysable observations could be obtained. Such a dataset was
collected at Petauke in 1954-60 on the incidence of rosette in relation
to time of sowing. This clearly showed that establishment percentages
decrease with delay in sowing, while percentage infection increased
enormously (Table 16.16). These two effects are confounded in this
analysis but it is known that establishment is less successful in delayed
sowings, quite independently of rosette infection. Observation also
suggests that poor gappy stands are more attractive to aphids, as are
chlorotic plants resulting either from virus infection or through waterlog-
ging. The suggestion has been made that, in closed stands, biological
control of the insect vector through entomophagous fungi controls virus
spread (which is less effective in poor stands). The attraction of aphids to
chlorotic plants draws the vector to the virus source in many instances,
from which it can spread rapidly. The rosette virus produces disease as a
rule in crops grown in poor conditions: the commonest factors producing
outbreaks are late sowing and drought. With good husbandry and III
seasons of good, well-distributed rainfall, it is not serious as a rule.
TABLE 16.16 Effect of time of sowing incidence of rosette, Petauke, 1959-60,
variety Mani Pintar
Sowing dates Total plants % plants No. plants % plants

infected infected harvested harvested
17 December 28 2.3 1218 81.2

31 December 76 6.3 1197 79.8
14 January 327 35.5 920 61.3
28 January 358 46.7 767 51.1
8 February 325 85.1 382 25.5
SE ±35 LSD P = 0.05 105
This is not the case, however, with leaf spot diseases. These are universal
and occur in good seasons and bad. They can produce very high levels of
yield loss in conditions particularly favourable for disease development.
The spanish/valencia group of varieties (subsp. Jastigiata) are much more
susceptible than are the virginia cultivars (subsp. hypogaea). Leafspot
control trials were carried out at Mount Makulu between 1957 and 1960
and results are presented in Tables 16.17 and 16.18. In the susceptible
Spanish 809 variety, yield reductions of >50% were produced in the first
season (1957-58) and c. 30% in the second (1958-59), in which the overall
yields of all treatments were higher. In the final season a comparative pair
of trials using Spanish 809 and the more resistant/tolerant Makulu Red
showed yield levels which were comparable in the fungicide treatments of
both trials; however, the control treatment in Makulu Red yielded
approximately 50% higher and still retained leaf at harvest, whereas
Spanish 809 was completely defoliated. There was also an interesting effect
on quality. In Makulu Red, kernel size was not significantly affected by the
disease; in Spanish 809, on the other hand, the proportion of larger kernels
doubled in the spray treatments. These experiments were conducted on
sites adjacent to areas in which the crop had been cultivated in the previous
season and a very high inoculum potential was ensured. Although no
investigations were carried out on possible means of delaying the onset of
leaf spot attack by the use of windbreaks and maximizing distances be-
tween sites of successive crops, these were thought to be areas worth
exploring as possible methods of cultural control.
There were other pests and diseases of sporadic occurrence which were
noted but not investigated in detail. Insects which caused some damage
included thrips, leafhoppers, caterpillars, white grubs and termites. No
control was attempted and problems never got seriously out of hand. In
retrospect it is perhaps as well that control by pesticides was not attempted
- natural control of these pests seemed quite effective with reasonably
stable pest populations
TABLE 16.17 Leaf spot control trials with fungicides, 1957-9 (mean yields,
kernels lblacre), variety Spanish 809
Variety: Spanish 809
Season: 1957-8 1958-9 mean 1957-9

Fungicide treatment (SE ±94) (±103)
Control 657 1486 1072

Sulphur dust 961 2105 1533
Bordeaux mixture 1566 2235 1901
Zineb 1391 2248 1820
Maneb 1399 2227 1813
LSD P = 0.05 286 311
TABLE 16.18 Effect of controlling leaf spot by fungicides on yield and quality of
two groundnut varieties at Mount Makulu, 1959-60
Variety: Spanish 809 Makulu Red
Fungicide treatment Mean yield % retained by Mean yield % retained by

kernels 19/64" x 314" kernels 19/64" x 314"
lb/acre screen lb/acre screen
(SE ±177) (SE ±182)
Control 1202 30.66 1822 91.55

Dispersible sulphur 2273 65.62 2329 93.04
Copper oxychloride 2193 61.73 2062 93.90
Zineb 2261 65.28 1923 92.53
Maneb 2070 56.98 2086 92.04
LSD P = 0.05 546 559
Diseases other than cercospora leaf spots and rosette were never serious.
Phyllosticta, Pleospora and Pseudoplea were sometimes isolated from leaf
lesions but attacks by these fungi never reached epidemic proportions.
Bacterial wilt (Pseudomonas solanacearum) was sporadic in occurrence,
often most severe in the first years after ploughing up resting land and
declining thereafter. Sclerotium rolfsii, although destructive where infec-
tion occurred, was only sporadic and effective disposal of crop residues by
turning in prevented this potentially serious pathogen becoming a prob-
lem. Seedling rots produced by fungi such as Aspergillus niger were
effectively controlled by seed dressings. Concealed damage of kernels
could be incited by a variety of species, including Corticium solani in stored
pods, but the local practice of shelling groundnuts as soon as practicable
after harvest reduced this problem.
TABLE 16.19 Groundut yields and cultural practices used by the winning competi-
tor in the Rhodesian National Groundnut Competition, 1972-74
1972/73 1973/74
Area of groundnuts 6 ha 40ha

grown:
Yield of 4 ha competition 9400 kg/ha 9600 kg/ha
plot (kgs/ha unshelled):
Other crops grown: Maize 94 ha Maize 50 ha Winter
wheat 25 ha
Variety grown: Makulu Red Makulu Red
Seeding rate: 150--160 kg/ha 140 kg/ha
Row spacing: 4 rows x 35 cm on 1.8 m 6 rows x 23 cm on
bed 1.8m bed
Fertilizer application: 400 kg/ha compound S 500 kg/ha compound S
6%:N
18%:P20S
6%:K 2 0
220 kg/ha gypsum 150 kgs/ha gypsum at 6
broadcast 8 weeks after and 12 weeks after
sowing sowing
Herbicide Dyanap Dyanap, alachlor
Foliage insecticide: None Nuvacron (20/11/73)
Leaf spot control: None Agricura special
fungicide plus dithane
applied 15/11/73,
15/12/73 and 17/1/74
Sowing date: 7 November 1972 1 October 1973
Digging date: 15 April 1973 1 March 1974
Growing season length: 160 days 150 days
Winner 1972173 and 1973174: B. Huntsman Williams Altitude: 1233 m

Farm: Portwe Estates Soil type: sandy clay loam
(d) Adaptive research conclusions

As a result of this experimental programme carried out between 1954 and
1961 (Smartt, 1967)) a package was produced for the grower which in-
cluded a high-yielding variety and a set of recommended cultural practices
to produce considerably improved levels of yield without involving addi-
tional inputs in the way of fertilizers, herbicides, insecticides or fungicides.
It was interesting to note some 11 or 12 years later in Zimbabwe that this
package, produced with the indigenous farmer in mind, was elaborated
very successfully into a highly productive system which involved high levels
of inputs. Details of the system are presented in Table 16.19.
The system itself has a number of interesting features. The crop was
sown early and irrigated until the rainy season set in. In the first season
only fertilizers and herbicides were used; in the second, three fungicide
sprays were given at monthly intervals and a single foliage protectant
spray. The result was pod yields in two successive seasons of 9.4 and
9.6 t/pods/ha (6-7 tlha shelled). The factors which produced this remark-
able yield were the availability of irrigation water and substantial avoid-
ance and/or control of leaf spot disease. Very high plant population
densities were established, with adequate water supplies to produce a fully
mature crop. The leaf canopy which produced this crop had been protected
from insect and fungus attack when necessary. It is interesting to note that
this was necessary in the second season but not the first. Data from further
crops produced under similar conditions in the same area would have been
of interest. With extension of the cultivated area in the two seasons, it
might be expected that leaf spot control would be necessary to maintain
yield level; if control of foliage-feeding insects proved to be necessary
again, it would suggest that perhaps natural population control mechan-
isms of the pest had been adversely affected. In this development of the
initial 1961 package, the two major constraints identified had been
removed: water availability limiting plant populations and the yield re-
duction due to leaf spot.
This indicates the need for clearly identifying the major yield constraints
so that they may be addressed progressively as and when this is practicable.
In the original 1961 package, the constraints were imposed by limiting
rainfall and the impracticability of leaf spot control by fungicides and of
weed control other than by mechanical means. In the 1972-74 develop-
ment, all these constraints had been removed by supplementary irrigation
and use of fungicides and herbicides. Experimental programmes should be
designed to answer basic questions and, in applying the findings, due
regard must be paid to practical considerations.
In designing such an experimental programme, it is desirable to make
the experiments as simple and straightforward as possible and to design
them to answer clear and well-defined questions. This is especially so with
experiments that are being conducted by research stations in farmers'
fields. These can be remarkably successful if the willing co-operation and
interest of the farmer is secured - and not only of the farmer but also that
of the extension service. Some of the most successful experiments carried
out in Zambia in the period 1954--61 were those designed at the Mount
Makulu Central Agricultural Research Station planted by extension staff
of the Southern Province on the family holding of a local Tonga farmer. All
concerned derived the greatest satisfaction from their successful outcome.
16.3.2 Adaptive research and agricultural development strategies

It has come to be appreciated that there is a very considerable gap between
the generation of research findings and their application to practical agri-
culture. This has led to the formulation of an approach to research aimed
at bridging the gap under the name of adaptive research - a new name but
not a new activity, as has been shown. It was practised in the later phases of
British colonial rule with some success (Smartt, 1967) but it has been in a
state of eclipse because of the success of the Green Revolution. Many of us
who had been working in agriculture in the developing world were hear-
tened by the great leap forward of this revolutionary change. Although its
shortcomings were recognized at a surprisingly early date (Brown, 1970),
to some it showed that the agriculture of the Third World could be
developed in a positive and constructive way and that wise and judicious
investment could be justified in strict economic terms. Prior to this, agricul-
tural development had been regarded more or less as an economic black
hole.
Possibly the most unfortunate consequence of the Green Revolution
approach was that it led the 'top down' strategy of development. The
assumption on which it was based was that improved practices adopted by
the more successful and wealthier farmers who could afford the new
technology would tend to trickle down to the lower levels of agricultural
society. This arose from the attempt to economize on the Green
Revolution package. Recommended agricultural practices, miracle seeds
and appropriate technology were transferred without the economic under-
pinning. Circulating loan funds were not made available,with the result
that those poorer farmers who attempted to invest in the new style farming
fell prey to money-lenders. From being small-scale peasant farmers, they
often became landless labourers.
In agricultural terms, the Green Revolution concept did not take any
account of traditional agricultural practices. It was a system of agriculture
designed on purely logical, scientific lines - in fact rather narrowly based,
with the emphasis on an area's staple crop (actual or potential). The
narrowness of the basic concept had a number of unfortunate conse-
quences in that improved high-yielding new cultivars made landraces of not
only the staple crop species but also those of other crops uncompetitive.
This tended to produce large areas of monocrops - conditions favourable
for the generation of pest and disease epidemics. A reaction to this, which
has set the clock back approximately 40 years, has been a return to systems
closer to traditional farming which tend to maintain stability of production
rather than the boom-and-bust cycles of monocrop agriculture that involve
large areas of genetically uniform populations of a single species. This
reaction has led to a reversal of strategy in favour of a grassroots-upwards
approach, involving close collaboration between research and extension
workers on the one hand and farmers on the other. It necessitates building
up knowledge of local farming systems and agricultural practices so that
the constraints in the system can be identified and defined and research
directed to their relief. In this way it is to be hoped that much (if not all) of
the inherent stability of the traditional system can be preserved while
productivity is improved.
It is obvious that the personnel involved in institutionalized basic re-
search do not include those who would engage most effectively in adaptive
research. Quite properly, the scientific staff at an international agricultural
research institute includes highly trained and specialized scientists, who
would not be expected to have agronomic training to any great extent. The
scientist engaged in adaptive research would need to have a good, broad
agronomic training and be able to collaborate effectively with specialists in
plant breeding, plant pathology, economic entomology and soil science. In
order to do so, such a scientist must have some training in all these fields,
even if only at a rather basic level. It is interesting to consider how, in the
past, such training was given. In the days of the British Colonial
Agricultural Service, there were two types of graduate who were recruited
for training: those with a training in agriculture on the one hand (who were
largely destined for extension or agronomic investigation) and, on the
other, graduates in the natural sciences who were trained as plant
breeders, pathologists, entomologists and soil scientists. The latter also
received a broad general training in agriculture and were able to appreciate
their own research field in its proper context. In addition, such individuals
were quite well placed to carry out experimentatioQ in fields related to
their own specialization in the absence (all too frequent!) of the appropri-
ate specialist. In the dependent territories 40 years ago, it was not uncom-
mon for pathologists to be instigating and executing variety trials or for
plant breeders to be carrying out agronomic investigations. With a broad
agricultural background and an adequate training in field experimentation,
this often worked quite well.
Another interesting approach has been developed at Wageningen, in the
Netherlands, where specialist training in plant taxonomy has been com-
bined with agronomy. This has produced a number of excellent agricultural
botanists who have made very significant contributions in the general field
of the botany and taxonomy of crop plants and especially in the specialist
field of crop plant genetic resources. At first sight this combination of
disciplines seems unusual, if not bizarre, but it shows a deep appreciation
of the kind of scientific personnel needed to maintain progress in research
and development for the Third World. There is a very real need for
versatility and adaptability in research personnel engaged in crop improve-
ment and agricultural development. In addition to the training and quali-
ties already mentioned, they need to be open-minded, receptive to new
ideas, very resourceful and able to carry out sound experimentation with
the simplest equipment. Fortunately simple experiments are often all that
are necessary. Complex experiments have a nasty habit of generating too
many missing plots for effective analysis, while simple randomized block
and Latin Square trials can prove to be remarkably resilient.
In colonial times, when agricultural research was undertaken by the
departments of agriculture in the various dependent territories, the quality
of research management was variable. In some instances it was so poor as
to inhibit any useful work being done; in others it was surprisingly good
due largely to commonsense combined with a basic appreciation of what
research was about and some inkling as to how it might be undertaken. All
too often the attempt was made with inadequate personnel. For example, a
single plant breeder might be made responsible for the improvement of the
full range of crops in the local agricultural calendar. Needless to say in such
conditions the effort was so widely dispersed and thinly spread as to be
totally ineffective. A concentrated but well focused effort on a narrow,
well-defined range of crops or even a single crop was much more effective,
provided of course that clear goals were set and that adequate logistical
support was available. Such efforts can be extremely productive and highly
cost-effective as long as appropriate exploratory work has been carried
out, the problems clearly defined and adequate procedures devised to
implement the research programme designed to provide solutions.
Research management implies more than the mere management of
research personnel. The current idea that a 'good manager' can manage
anything from a whelk stall to a giant multinational enterprise can be very
far indeed from the truth. The manager must know and understand what it
is that is being managed. The ideal manager would be a polymath with a
good background in and understanding of the earth and physical sciences,
in addition to considerable experience in the biological sciences as applied
in the agricultural context. An ability to communicate with agricultural
economists and sociologists would be advantageous. Such paragons do not
exist in any great numbers! Perhaps it is more reasonable to expect to find a
broad-minded agricultural scientist who can look inwards to the work of his
research organization, organize it in a coherent and integrated fashion and
not only package, present and 'sell' its product but also be fully cognizant
of the needs and aspiration of those outside the organization, especially
those whose activities have interfaces in common.
16.4 CONCLUSIONS
In any research endeavour, it is important that both the immediate and the
more remote contexts of the work be fully appreciated. That is why
farming systems and adaptive research have been considered at some
length in this chapter. In specialist crop circles it is very easy indeed to lose
sight of such considerations. The awareness of the relationship between
one's own work and that of one's colleagues greatly facilitates the flow of
information. Information can be one of the most underused of our re-
sources - not least that which, although gleaned decades ago, remains of
considerable current relevance. Lack of awareness of previous work leads
to attempts to re-invent the metaphorical wheel or complete failure to
learn the lessons of the past. We in the groundnut world have the spectre of
the East African Groundnut Scheme to remind us of the consequences of
References 699
ill advised and poorly considered development projects. The present era is
that of the 'instant fix' the politician is constitutionally impatient and tends
to favour the short-term 'solution' to development problems produced by a
consultant, notwithstanding the totally inadequate factual base on which
developmental initiatives might be founded. Quite obviously resources
would be better directed into a programme of on-the-spot adaptive re-
search, which could ensure that development could be both compatible
with the local farming systems and sustainable in the long term.
REFERENCES
Allan. W. (1967) The African husbandman. Oliver and Boyd. London.

American Society of Agronomy (1976. reprinted 1977) Multiple Cropping. ASA Special
Publication no. 27. Madison. Wisconsin.
Brown. L.R. (1970) Seeds of change. Praeger. New York.
Evans. A.e. (1954) A study of a rosette resistant variety Asiriya Mwitunde. East A/i"ican
Agricultural Journal. 22. 27-31.
Hammons. R.O. (1973) Genetics of Arachis hypogaea. in Peanuts - culture and uses.
American Peanut Research and Education Association. Inc .. Yoakum. Texas.
Hartley. W. (1949) Plan! col/ecting expedition to sub-tropical South America. Report Division
of Plant Industry Australia no. 7 pp. 96.
Hildebrand. G.L. and Smartt. J. (1980) The utilization of Bolivian groundnut (Arachis
hypogaea L.) germ plasm in Central Africa. Zimbabwe Journal of Agricultural Research.
18.39-48.
Loomis. R.S. and Connor. D.J. (1992) Crop Ecology. Cambridge University Press.
McEwen. J. (1961) Groundnut variety Man! Pintar. Nature. 192.92.
Ruthenberg. H. (1976 and 1980) Farming systems in the tropics. 2nd and 3rd edns. Oxford
University Press.
Smartt. J. (1960) Genetic instability and outcrossing in the groundnut variety Man! Pintar.
Nature. 186. 1070-1071.
Smartt. J. (1967) Groundnut production in Zambia. Government Printer. Lusaka.
Smartt. J. (1978) Makulu Red - a 'Green Revolution' groun<.lnut variety. Euphytica. 27.
665-675.
Tivey. J. (1990) Agricultural Ecology. Longman. London.
Trapnell. e.G. (1953) The soils, vegetation and agriculture of North-eastern Rhodesia.
Government Printer. Lusaka.
Trapnell, e.G. and Clothier. J.N. (1957) The soils, vegetation and agricultural systems of
North-western Rhodesia. Government Printer. Lusaka.
Vandermeer. J. C. (1989) The ecology of intercropping. Cambridge University Press.
CHAPTER 17
The future of the groundnut crop

]. Smartt
In assessing the future prospects of any crop, it is sensible to consider what

factors contribute to the crop's present position and how these are likely to
change, if at all, in the immediate and near future. The considerations tend
predominantly to be economic in that the output from producing the crop
must be commensurate with the input, though the input may not always be
easy to quantify in purely monetary terms. Economic viability needs to be
assessed by a broad and open-minded economist, not by an accountant
who will tend to ignore what cannot be expressed in cash terms.
17.1 PRODUCT DIVERSITY
There are crops such as coconuts and bananas for which the level of inputs
(in some circumstances at least) may be quite low. After the crop is
established the farmer has little to do but to watch it grow and harvest it. A
little gentle weeding and cultivation may be all that is needed. This is never
the case with groundnuts: the returns, good though they may be, are not
won easily nor without considerable effort. One might ask at this stage why
the effort still continues to be forthcoming and the reason lies in the very
high regard in which the product is held and its great versatility in use. The
uses vary quite remarkably in the various areas of production. In subsis-
tence economies the seed is used directly for human consumption while the
crop residue is a useful livestock food. It is interesting to note the use that
was made of the crop in the United States for 'hogging off', in which
groundnut crops at or near maturity w'ere in essence harvested by pigs. It is
not beyond the bounds of possibility that such usage, given favourable
circumstances, might return.
The utilization of the groundnut as food by subsistence cultivators is
varied -limited, one might say, only by human ingenuity. The crop can be
Product diversity 701
used in similar ways to peas and beans in what might be called the 'green
mature' stage before the fruit dries out, or even earlier when the pod is soft
and juicy (Krapovickas, 1969). According to the same author, it can be
used as the source of a refreshing drink, chicha de mani, and also a soap
can be made from the seed by saponification. Since the seed may contain
50% of its dry matter as oil, it is a relatively simple matter to extract the oil.
The mature seed can be parched or ground to a paste and used in the
preparation of soups and relishes. The seed is also amenable to use in the
traditional fermentation technologies practised at village level in south-east
Asia and the Far East.
By the nature of things, subsistence cultivators cannot be totally self-
sufficient: they usually produce some surplus which can be sold or
exchanged locally for other essential goods. There is usually some small
local trade in the crop since it is such a popular article of consumption. This
surplus provides the base from which (in favourable areas) an export trade
might develop. Historically, with urbanization there was no doubt a
demand from town and city markets for this popular and well-regarded
food material. The overseas export market was initially for the oil: Archer
(1853) reports that the crop was sold by dry-fruiterers but the bulk of
imports went to the oil mills. He noted, however, that the supply was
irregular - 'sometimes several hundred quarters in the year, and at other
times none at all'. He went on to say: 'It is never likely to be much used as
an edible fruit in this country [Britain], being in no respect superior to the
common grey pea of our fields.' Clearly the development of export markets
depended on the reliability of production and relative stability of markets
which has developed in the past 100-150 years. Johnson (1964) presents an
extensive historical review of the history and past utilization of the crop
and shows how, in the United States, slow progress was achieved towards
general acceptance of the groundnut by the apostle of the peanut, George
Washington Carver, in the early part of the present century.
The bulk of initial exports found an industrial market in the edible oil
expression industry. The press cake became a valuable and lucrative by-
product and a well-regarded, high-protein feeding stuff, particularly for
dairy cattle, in the first half of the present century. In North America, the
edible oil industry was eclipsed as a market with the development of
confectionery uses in candies and sweets in addition to traditional roasted
and salted peanuts, but most of all by the development of the peanut butter
market. Peanut butter, together with cola beverages, has become some-
thing of a North American addiction.
The result of all this is that the groundnut has an enormous diversity of
use at present, which tends to promote stability of markets in the long
term. It does mean that the quality prescriptions of the market must be
met. Then there is the question of adjustment to changes in market
demand. That for groundnut as an oilseed is slackening for a number of
reasons, principally because of competition from other oilseeds (maize,
702 The future of the groundnut crop
sunflower, soya and rape) but also because of a change in the nature of the
material exported. Until comparatively recently, oil expression depended
on imported stock but there has been a tendency for this process to be
carried out in the country of production. Unfortunately this has led to the
problem of adulteration of groundnut oil with cheaper oils and, naturally,
the reputation and standing of the product has suffered.
In contrast, the demand at the upper market level has remained good
and is buoyant. This area is particularly attractive to food and confec-
tionery manufacturers because the added value potential is enormous. The
retail value of a tonne of groundnuts sold as roasted peanuts (approxi-
mately 40000 packets) compares very well indeed with the import cost of
the raw material. Just as there has been an attempt by producers to reap
some benefit from groundnut processing in production of oil and press-
cake, so it is reasonable to expect that they may well wish to benefit from
the added value arising from peanut butter and confectionery manu-
facture. This could be attractive if the groundnut hulls could be used as an
energy source for manufacture. The groundnut confectionery sector might
well seek to emulate the processing of sugarcane as an eminently energy
efficient industry.
Meeting the market demands for such an industry poses a challenge and
imposes some constraints. Quality standards for oil stock have not been
high - in the United States only reject nuts are used and the problem of
aflatoxin is less important as it remains in the cake and is virtually absent
from the oil. For confectionery use, quality standards are very exacting
indeed: aflatoxin levels must be low and demands for even lower levels are
likely to be imposed. The physical properties of the kernels are important;
size, shape and even testa colour (although removed in blanching) must
conform to standards. Chemical and organoleptic properties are similarly
important.
The multiplicity of outlets and the diversity in utilization provides stab-
ility in the market, which is a great advantage in planning future pro-
duction initiatives since these can be based on a firmer foundation than
might otherwise be the case. However, the market's exacting standards
must be met and fully taken into account in the course of planning.
17.2 FARMING GROUNDNUT
The groundnut is a high-input crop at all levels of production. It is basically

demanding in terms of labour and time for land preparation, sowing,
cultivation, harvesting, stripping and shelling. The time and labour com-
mitments can be reduced by mechanization and the use of chemicals to
reduce cultivation. In more advanced production systems, applications of
gypsum or land plaster and insecticides and fungicides are commonplace
Farming groundnut 703
and routine. These inputs are additional to those used by the less sophisti-
cated cultivator.
17.2.1 Levels of input and productivity

In Central Africa it used to be a rule of thumb that a yield level of about
200 kg/ha was critical. Below this level the subsistence cultivator would not
find it worthwhile to produce the crop; above, it was worth considering.
The level of input necessary for a shifting cultivator was the effort required
to clear and prepare the land, sow and cultivate the crop, harvest it and
process it post-harvest. The inputs could be considered as labour, time,
seed and the use of simple tools such as the hoe. With increasing sophisti-
cation the farmer in the developing world, operating above the subsistence
level, will be able to extend the cropping area if animal power is available.
The use of animal power incurs the need to invest additionally in at least a
plough, and probably a cart and cultivating machinery. While the farmer
may be relatively self-sufficient for animal power, clearly the operation
must be capable of generating a cash surplus to pay for the capital cost of
equipment. This is even more the case when mechanization takes place: a
further reduction in self-sufficiency is incurred as well as expenditure on
fuels, lubricants and maintenance costs in addition to capital and interest
payments. This generates further pressure to increase either the land area
cultivated or its productivity, or both. At the limit of available land area,
productivity increases are necessary in order to meet these additional costs.
Productivity can be improved by the application of fertilizers, and by the
use of pesticides (herbicides, insecticides and fungicides) to control losses
due to pest depredations. The increased costs must be scrupulously
balanced against additional income produced and the rising curve of
increasing expenditure must be less steep than that of income. Obvious
though this is, it can be overlooked - especially when the farmer, as in the
developing world, does not have the services of an accountant giving
advice about what is producing a sufficient return and what is not.
17.2.2 Energy budgets

An area of increasing concern is that of the energy budgets of different
production systems. In terms of productivity per unit of non-solar energy
input, it has been claimed that the least sophisticated systems are the most
efficient and the more sophisticated the least so. The energy investment
from outside the farming system begins with the use of the hoe, the
manufacture of which involves comparatively small amounts of energy.
This increases with the use of ploughs, cultivators and other tools
that require animal power. Additional increases are incurred with
mechanization, the consumption of oil fuels, the manufacture of agri-
chemicals and the energy cost of their distribution. In the most advanced
systems of groundnut production, where field-curing is abandoned, there is
considerable energy expenditure in drying the crop. The sum total of
energy expenditure in producing the final, saleable crop is substantial.
The point can be made, with some justification, that the energy which is
expended in the production of the crop may, if the trend continues, be
greater than th~ intrinsic useful harvested energy in the crop itself. Such an
energy deficit would dearly be made up by using fossil fuels. This poten-
tially negative energy balance is a question of which we would be aware
and attempt to address with circumspection. The question can be asked:
how are we to set about improving the energy balance equation? We need
in effect to be energy misers, using materials such as fertilizers only insofar
as they give a sensible return and applying them at the most opportune
times. In addition we should be seeking to reduce applications of pesticides
by increasing levels of pest and disease resistance, seeking energy efficient
use of herbicides (or alternatives to them) and the use of natural energy as
far as possible in drying the crop. The energy balance may, in spite of
improvements, remain adverse. Another pressure to which the producer
is subjected arises from the pest residue problem. Economy in the use of
agrichemicals may thus be doubly friendly to the environment.
17.2.3 Aflatoxin
The aflatoxin problem, which has been recognized for 30 years and more
(Chapter 13), has not been finally resolved, although there has been
progress in knowledge of the causal organisms and the way they produce
their effects. Possibly the most productive approach is first of all to attempt
known environmental approaches to minimize invasion, then to select host
genotypes in which penetration of the pod wall is slow and finally to select
host genotypes which do not induce mycotoxin production by the fungi.
The prospects of more rigorous standards and lower tolerance levels is a
very real one. Conditions of storage in producing countries also need to be
of a high standard so that produce· is not denied entry into importing
countries. Continued mycological and breeding studies are required.
17.2.4 Climate changes

Climate change and the spread of cultivation to new areas, most notably
south-east Asia, both raise problems. With shorter and less reliable wet
seasons in some areas, particularly in sub-Saharan Africa, there is con-
siderable interest in the breeding of ever more early-maturing cultivars.
The questions are: what is the shortest period to maturity in which a useful
and worthwhile yield can be achieved, and what is the minimum canopy
Farming groundnut 705
size with which this end can be achieved? Success has been achieved with
ultra-early maturing cowpeas and this could be repeated in the groundnut.
Something like the valencia growth form might be appropriate with a
pattern of assimilate partitioning producing a high harvest index.
17.2.5 Cultivation in new areas: adaptive research

In south-east Asia the problem is quite different. The groundnut in the
tropics has been a feature of semi-arid environments where a single crop is
taken per year. In the more humid tropics, there is a problem which arises
from the fact that it may be possible to grow the crop all year round: the
question then is to determine how best to go about this. A difficulty can
arise in drying seed post-harvest, which is important for maintaining seed
stocks. No problem arises for that part of the crop which is boiled for
eating but for the part of the crop which is to be dried the drying process
must be continuously maintained. Similar procedures to those developed
for drying cocoa beans might be suitable. In areas where convectional
rainfall is characteristic, the drying crop could simply be protected at the
appropriate time of day. Periods of unremitting rainfall, however, could
cause difficulties with moulding and mycotoxin generation.
The problems which could arise from effectively continuous groundnut
cultivation in the humid tropics need to be evaluated very carefully.
Certainly in the ~emi-arid tropics the occurrence of a close season has been
regarded as an advantage. If cultivars with short-term seed dormancy are
grown, then there are few (if any) volunteer plants to carryover viruses,
vectors and fungal pathogens directly between successive crops. There
would be no such check with continuous cultivation and the consequences
of this need to be studied. These are not necessarily what might be
anticipated. In areas where three and perhaps even four crops might be
taken in a single calendar year, it is quite possible that one particular time
of year might have particular advantages. This could be advantageous vis a
vis the position in the semi-arid tropics (S.A.T.) where debris from one
season's crop persists through the dry season and produces inoculum for
the cercospora leaf spots at the beginning of the following rainy season. In
the humid tropics were there to be a close season the debris would have
decomposed before the next crop was sown.
While the agronomic position is well explored in traditional areas of
groundnut cultivation, current studies are regarded as fine tuning. There is
scope for and a need to explore the situation thoroughly in areas where the
crop has been introduced comparatively recently. The question of cultivar
selection is obviously an open one, too. For example, is it more productive
to produce four crops per annum of short-season cultivars than three of
long-season varieties? It would be worthwhile to explore the performance
characteristics of the very longest season varieties and it would be useful if
comparable results could be obtained from a single long-season crop as
from two crops from an early maturing cultivar. There is a supposition that
late-maturing forms tend to have low harvest indexes and would not be
viewed favourably if seed production was the major concern. However, if
the haulm was produced in a useful amount and was of good quality, such
varieties could have a useful role. In such novel situations it is essential to
be both objective and open-minded: received ideas from elsewhere may be
of very limited application. The basic knowledge we have of the crop
should be the starting point for a programme of adaptive research. Such a
programme should take into account the objectives which are practicable
in the light of present constraints but one should also bear in mind other
objectives which might be explored when present constraints no longer
operate. While we must obviously take into account present realities, we
should keep a weather eye open for future possibilities.
17.2.6 Nitrogen fixation

The great advantage claimed for legumes in general (and not always
realized) is that they are virtually self-sufficient for nitrogen. In the semi-
arid tropics, this is very largely true. At the end of the dry season the first
rains initiate a flush of nitrification by free-living soil bacteria which early
sown crops are in a position to exploit. An early-sown groundnut crop in
such circumstances can make use of this available soil nitrogen while its
own nodular system is built up. With further rainfall, soil N levels can be
expected to decline as the nodules on groundnut roots become effective.
Later-sown crops do not derive any such benefit and may suffer not only
the disadvantage of less soil nitrogen but also soils which are cooler and
less well aerated on account of the rain. The crops establish more poorly as
a result and they are also more prone to infection by aphid-borne viruses
and to receiving leaf spot inoculum very early in their growth.
Root nodules in the groundnut are formed by Bradyrhizobium (Chapter
8) and there has been considerable research into improvement of the
efficiency of N2 fixation in the root nodules. Much attention has been
focused on selection of strains which are especially efficient in N2 fixation,
in the hope that these might be released and establish themselves, infect
roots and produce more effective nodules. This strategy appears to fail on
two counts: that laboratory selected strains are often less competitive in the
wild than the native bacteria, and that there is no possibility of establishing
super-efficient Arachis-specific Bradyrhizobium populations because the
cowpea cross inoculation group is all but universal in the tropics and
subtropics. The best approach (Chapter 8) seems to be to select host
genotypes with a propensity to produce effective nitrogen-fixing nodules
with a very broad cross-section of the native Bradyrhizobium population.
It is possible that the nitrification system might be manipulated in this way
to advantage. Selection and evaluation would probably not be straight-
forward, however.
Farming ground nut 707
17.2.7 Fertilization
Fertilizer studies in the groundnut are fraught with difficulty. There is no
question that the crop, judged by the yields it produces, can respond
handsomely to high fertility conditions. The experience of agronomists
who have attempted to study fertilizer responses has frequently been one
of frustration, the exception being calcium response. On calcium-deficient
soils there are clear-cut responses to applied gypsum or lime in terms of
improved seed development and reduction of 'pops'. Apparent response to
application of other nutrients is erratic and the reason seems to be that the
nutrients are not in fact reaching the soil horizon in which groundnut roots
are actually taking them up. Deep placement of nutrients is not practicable
and the received wisdom of 40 years ago (York and Colwell, 1951) is
probably as valid today as then: fertilize the rotation rather than the
groundnuts. They will pick up nutrients which have passed beneath the
root range of other crops and will not be wasted. This is consistent with the
observed calcium response: the pegs and pods which take up this element
are in the top 5 cm of soil and top dressings are readily accessible. The
problem of apparent lack of response is actually one of accessibility, and
incidentally probably led to the soubriquet of 'the unpredictable legume'
being applied to the groundnut. It is less unpredictable now than it was,
perhaps, but it is still by no means an open book.
17.2.8 Mixed cropping

The role of groundnuts in farming systems, particularly those appropriate
to the developing world, would probably repay further study. The rela-
tively low incidence of pests and diseases in traditional farming practices
has often been remarked. In Mexico (Miranda, personal communication),
the Mexican leaf beetle became a serious pest when the Phaseolus bean
ceased to be grown as a mixed crop with maize and was cultivated as a sole
crop. Mixtures of crops and mixtures of genotypes within a crop stand can
both exert a controlling effect on pest and disease incidence. The challenge
is how to gain advantage from the lower pest and disease incidence in
mixed crops and yet benefit from mechanization and advanced technology.
Basically similar approaches are adopted in relay sowing and in agro-
forestry. The effects of combining species of high and low growth can have
beneficial effects in delaying arrival of inoculum and reducing spread by
wind. I have observed instances of groundnut crops grown in relatively
small woodland clearings which had markedly lower incidence of leaf spot
diseases. It seems that anything which breaks up the uniformity of the
potential host substratum can have a beneficial effect in reducing pest and
disease incidence.
Such systems of mixed cropping have other beneficial effects in that the
soil becomes comprehensively permeated by root systems of different
species exploiting different soil horizons. This results in more efficient
exploitation of the soil's bank of nutrients and recycling them and at the
same time more effectively binding the soil and providing protection
against erosion. There is wide scope here for the study of relay cropping
systems and agroforestry in conjunction with local traditional farming
systems, with a view to identifying the constraints which inhibit their
further development and removing them to enable development to
proceed.
17.2.9 Water resources

The question of water resources and their effective use is another import-
ant issue which has to be addressed. Groundnuts have a reputation as a
drought tolerant crop. They owe this mainly to their deep-rooting habit.
While they may survive drought conditions if these are not too prolonged,
they may not produce a satisfactory yield. However, their tolerance of
drought can be put to good use in the semi-arid tropics. At the end of the
dry season it often happens that there is a false start to the rains and there
may be sufficient rainfall to allow preparation of land for sowing to be
completed. The natural sequence of events is to follow this with the sowing
of the staple cereal crop. In the case of maize, when no further rain falls for
a few weeks the crop may emerge and be lost from drought. Groundnuts
sown under such conditions will germinate, send down quite a long tap-
root, produce a small tuft of leaves and sit out the drought. Further growth
and development occur when the rains resume.
Groundnuts do require a good soil moisture status from flowering
through pod-filling. Shortage of water during this period results in poorly
filled pods and low quality kernels (Chapter 9). In terms of pod-filling,
plant population density must be related to expected soil moisture contents
at this critical stage. If it is inadequate, the level of yield may be maintained
but the quality of the product is drastically affected. Supplementary irriga-
tion (if water and the means to distribute it are available) could be used
quite effectively to increase yields of good quality. This would enable the
larger pod set from a high plant population to mature a larger crop more
fully. If the length of rainy season is limiting but irrigation water is
available at the end of it, groundnut crops could be started off under
irrigation and maintain themselves with relatively low water use until the
major rains come. This would give the benefits of a longer growing season
combined with the advantage of the crop having a longer period free from
pests and disease before pest and pathogen population densities build up.
This is an interesting area for further exploration by both the water-
relations physiologist and the irrigation specialist. In addition there is
scope for detailed study of growth and development of root systems, with a
number of ends in view, providing useful information for those investigat-
ing rhizobial activity and the uptake of both water and nutrients.
Improving the groundnut 709
Comparative studies of the patterns of root growth in different genotypes
and the morphological types of the groundnut would be particularly valu-
able, though not an easy area for the breeder to explore. The root system
of the groundnut underpins and supports so many of its useful attributes
that we really ought to know more about it.
17.3 IMPROVING THE GROUNDNUT
Since the pioneering days of Gregory, Smith and Yarborough (1951), there
has been a continuing development of botanical knowledge which provides
the foundation for much of the work of groundnut improvement. In many
areas the situation is highly satisfactory but in others there are large
lacunae which have to be filled. Our basic morphological and anatomical
knowledge of the cultigen and its wild relatives is reasonably satisfactory
(Chapter 3). Our knowledge of its reproductive biology and development
is also quite satisfactory (Chapter 5). This is not to imply that nothing
further remains to be done in these areas, but there are others where lack
of information and deficiencies of published knowledge are a decided
handicap, especially in biosystematics related to genetic resources (Chap-
ter 4). This has repercussions affecting the use of genetic resources in
breeding (Chapter 14) as well as the development of new uses for members
of the genus as forage in addition to their use as a genetic resource for
improvement of the cultigen (Chapter 15).
17.3.1 Taxonomy and biosystematics

Since the 1950s there has been an active interest in the collection of the
genetic resources not only of the cultigen Arachis hypogaea but also its wild
relatives. The volume of material collected by Krapovickas and Gregory
has been most impressive. Collection has been carried on in all areas of
South America where the genus is known to occur wild, and also of
landrace materials, particularly in the Americas and Africa. Unfortunately
the substantial volume of wild material collected (including many species
new to science) remains undescribed. Some of it has been given informal
names and an informal taxonomy has been put forward (Resslar, 1980).
This has been useful as far as it goes but it does not go far enough. The
users of wild species as genetic resources need to know the status of their
material to guide them in planning strategies for its use as a germplasm
resource and in putting their plans into effect. The reasons for the delay are
not clear but it is a matter of some urgency that this particular nettle is
grasped and that a scientifically sound monograph be published of the
genus as it is known at present. It is unrealistic to hope that a definitive
monograph can be produced now which will stand for ever without
revision - after all we must provide some useful future employment for
taxonomists! The field of taxonomy and biosystematics is a vital one and
essential to the development of sound and sensible strategies of genetic
resource utilization. It is possible that this particular programme of work is
under-resourced and that the first priority has been that the material in the
wild be collected before it is made extinct by encroaching development and
habitat destruction. The taxonomic synthesis may have had to take second
place. We need to know what the situation is so that any necessary
representations can be made and appropriate pressure applied where
necessary to secure funding for the conclusion of this very necessary work.
17.3.2 Evolutionary history

In order to make effective use of wild Arachis species as a genetic resource
we need to understand the evolutionary history of the groundnut itself
(Chapters 2 and 4). Interesting problems are raised by the fact that the
cultigen is a tetraploid which, together with its wild prototype Arachis
monticola, constitutes a single biological species and that it is placed in an
otherwise diploid section of the genus. The wild prototype material that is
available amounts to a very slender genetic resource but the diploid species
(most of which can be crossed with the cultigen) together constitute a very
valuable resource. In spite of the ploidy level difference, it is possible to
induce gene flow from them into the cultigen. Some species have been
identified as sources of pest and disease resistance, among them Arachis
cardenasii - one of the first species to produce viable and partially fertile
triploid hybrids with the cultigen. It has resistance to late leaf spot and
nematodes (Stalker, 1992) and is being used in breeding programmes at
present. Studies of interspecific hybrids between diploid species have
shown that three distinct genomes probably exist: A and B (Smartt et al.,
1978) and D (Stalker, 1991). It has been suggested that A and B genomes
are present in the cultigen. While similar in genetic complement, these
genomes are differentiated from each other by differences in chromosome
structure and this confers a high level of genetic stability in the amphi-
diploid (allotetraploid) cultigen. There have been attempts to identify
genome donor species and the leading candidates at the present time are
A. duranensis and A. batizocoi. This hypothesis is supported by the results
of experimental crosses involving the cultigen, wild species and experi-
mentally produced amphidiploids. The suggestion has also been advanced
(Singh, 1988) that the two groups virginia and spanish/valencia might have
arisen from separate amphidiploidizations. This idea is supported by the
fact that there does seem to be some chromosome structural differentiation
between the two subspecies.
While the hypothetical ancestry of A. hypogaea is supported by the
studies of Klozova et al. (1983a,b) it has not been supported by the RFLP
studies of Kochert and co-workers (Kochert et al., 1991). RFLP evidence
suggests that A. hyspogaea is monophyletic: results from all samples tested
show very little variation. Diploid species of section Arachis do show
variation. Assuming that RFLP is an additive character it is possible, by a
summation of the separate RFLP patterns of A. duranensis and A. ipaenis,
to produce a combined pattern similar to that of A. hypogaea. However,
both A duranensis and A. ipaensis are 'A' genome species and an amphi-
diploid produced from this pair of species would not be cytogenetically
stable. There are suggestions (Kochert, unpublished) that more than a
single RFLP pattern may occur in a species such as A. duranensis and it
may well be that further search is necessary to identify complementary
morphs from two species which would produce the A. hypogaea pattern
and be cytogenetically credible. It is important to determine the extent
to which gene exchange or recombination is possible between the A and B
genomes. Some quadrivalent formation is observed from time to time in
the meiosis of A. hypogaea, which could possibly bring about some interge-
nomic gene transfer. This could facilitate expression of desirable recessive
characters and enhance expression of additive genes.
17.3.3 Role of the plant breeder

Sooner or later, and inevitably, crop improvement involves the breeder.
We have already considered that the work of the breeder would be
appropriate in the reduction of susceptibility to aflatoxin contamination,
the improvement of the efficiency of rhizobial activity by host selection,
and the improvement of root growth and function, to which also can be
added that of the shoot in physiological terms. The range of breeding tasks
which can be assigned to the groundnut breeder is enormous.
(a) Genetic resistance

One of the most attractive areas in which the plant breeder can operate is
in the improvement of resistance to pests and disease attack. The high
levels of mammalian toxicity in many insecticides, and their persistence,
have aroused particular concern. There is also the fact that many have lost
their efficacy in a very short time and that they are expensive and are costly
to apply. Similar considerations apply to fungicides.
Successful resistance breeding provides both a cost-effective and a
'green' solution to this kind of problem. Even a partial solution has value:
if application of agrichemicals can be delayed and their frequency of
application reduced, this is of positive value. Pest and disease control by
this means could enhance profitability considerably. But the breeder must
not be complacent - there are instances where breeders have exacerbated
the problem they aimed to solve, as in the loss of horizontal resistance in
cereal rust breeding. The entomologist faces a similar situation in that the
use of insecticides on the cotton crop has exacerbated the insect pest
problems there.
The increasing levels of mammalian toxicity in insecticides and their
decreasing periods of effective use have given urgency to the search for
alternatives. This has led to the rise of integrated pest management (IPM) ,
which takes an ecological approach in seeking how best to manipulate pest
problems by maintaining their populations below damaging levels.
Genetically controlled resistance or tolerance can be a component of such
systems in combination with cultural control, biological control and mini-
mal use of agrichemicals.
Control of virus diseases by breeding is interesting. The East African
groundnut variety Asiriya Mwitunde was shown in the early 1950s to be a
very poor aphid host and although it could be readily infected with the
chlorotic rosette virus was field resistant (Smartt, unpublished). Genetic
resistance to the virus itself has been found in West Africa and has been
incorporated in many new varieties. Cultural control of rosette in good
growing conditions is very effective but yields can be reduced to almost
zero under conditions of drought and poor stands.
The leaf spots are the universal pathogens of the groundnut and are the
cause of American growers undertaking costly spraying programmes. High
levels of resistance to both early and late leaf spots would save a great deal
of money where spraying is practised and bring about considerable and
virtually cost-free yield increases where it is not. Wild Arachis species such
as A. cardenasii and' A. chacoensis have been shown to have leaf spot
resistance and breeding efforts have been concentrated on these.
However, as anyone who has worked with large numbers of accessions in a
germplasm nursery knows, there are very considerable differences in leaf
spot susceptibility between accessions of the cultigen. There is a distinct
possibility that we may be overlooking useful sources of resistance here.
Hemingway (1957) showed that there are morphological resistance mech-
anisms, and a study of a possible range of morphological and physiological
mechanisms in South American landrace material might be productive. If
such resistances exist, they might be combined in a single variety which
might very well have a considerably higher level of resistance than any
current variety.
Nematodes are a problem in many areas (Chapter 10) and there is
considerable interest in controlling them by genetic resistance. Chemical
control is difficult and expensive and thus impracticable on a large scale.
Genetic resistance has been identified in wild species and active attempts
are underway to transfer this to commercial cultivars. Pest and disease
controls by incorporating genetic resistance are in essence the means of
protecting potential crop yield in the field.
(b) Efficiency of physiology

There is a continuing study of the physiology underpinning the production
of high yields and the improvement of harvest index. Manipulation of the
type of canopy produced is possible by genetic means, as is the partitioning
of assimilates between the various sinks. Selection for an efficient physi-
ology is clearly a complex matter. It is unlikely that there will be a single
optimal idiotype but it may be that a series of 'horses for courses' idiotypes
might be devised which would enormously enhance the efficiency and
effectiveness of the exchange of breeding materials internationally. It is
quite possible that certain materials may prove to be in effect pre-adapted
to a number of regions other than that in which they were produced. A
good illustration of this is furnished by the strong and clear pre-adaptation
of Bolivian landraces to conditions in Central Africa (Smartt, 1978;
Hildebrand and Smartt, 1980).
(c) Nutritional value and chemical composition

There is potential for breeding and selection work in the improvement of
chemical composition and nutritional value, on which little has as yet been
done. For example, contents of polyunsaturates and susceptibility to oxi-
dative rancidity are of concern to processors. The high concentration of
polyunsaturates held to be desirable by nutritionists can be combined with
a high content of the antioxidant tocopherols, a means by which this
particular circle can be squared. There is some interest in the confectionary
trade in kernels with intrinsically lower oil contents and these could in all
probability be achieved by selection. However, the same genotype (such as
Chalimbana) may produce mature kernels of very different oil contents
when grown at different elevations in Central Africa - those at lower
elevations having higher oil contents than those from higher altitudes.
Protein content, protein quality and organoleptic properties are also
heritable and could be bred and selected should the need arise. Other
commercial specifications set by the trade concerning heritable characters
(such as size and shape of seed and pod, and testa colour) cannot be
ignored but often have rather unfortunate repercussions on the grower and
engender great frustrations for the breeder. This is well exemplified by the
case of the cultivar NC2 when it was nearing the end of its useful life as a
major cultivar in the peanut belt of Virginia and North Carolina. In the
1950s and 1960s it met the millers' and processors' needs perfectly; they
had no interest in changing varieties and had apparently set their faces
against such a move. Eventually the supply situation deteriorated so
drastically that they were forced to change. Market forces will prevail
ultimately but their operation should be tempered by intelligent antici-
pation and vision and a consensus between all sections of the industry.
Quite justifiably, the breeders felt aggrieved that new varieties were
rejected because the commercial specification was not met by only the
narrowest margins. The industry appears to have learned its lesson but at
considerable cost and damage to itself.
17.4 THE ROLE OF BIOTECHNOLOGY
In the opinion of many, the future of crop improvement seems to lie in the
hands of the biotechnologist. To many in the field, this sounds like an
enormous oversimplification. It would be equally ill-advised to deny the
possibility of potential positive value as it would be to entertain totally
unrealistic ideas about what can actually be accomplished that is of practi-
cal value - and the cost-effectiveness of attempting to do so (Nigam, 1992).
Wild extrapolations from present accomplishments can lead to the setting
of over-ambitious targets in the future and almost inevitable disappoint-
ment. In biotechnology, if we do not manage to keep our feet firmly on the
ground, we are likely to dress up flights of fancy as realistic expectations. In
other words,we shall re-run the time-honoured bandwagon saga yet again.
In this connection it is interesting to consider a bandwagon which was in
motion 40 years ago - that of mutation breeding, which involved one of the
giants of peanut research, Walton C. Gregory of North Carolina State
University. His studies of mutation breeding were classic and well worth
carrying out for the light they shed on the operation of the genetic system
of the groundnut in particular and crop plants in general. In terms of useful
output of cultivars released, NC4X resulted but not a great deal besides.
New cultivars could be produced but less efficiently than by conventional
breeding programmes. There is no reason why useful mutations artificially
induced should not be fed into conventional breeding programmes. Where
mutation breeding has made its mark and continues to be of high value and
significance is in the improvement of vegetatively propagated crops.
Artificial mutagenesis plus selection of somaclonal variants have revolutio-
nized the breeding of chrysanthemums. It is a highly cost-effective breed-
ing technique for such decorative species and produces results in a much
shorter time than traditional breeding methods.
There are a number of reasons why the potential value of biotechnology
in groundnut improvement should be assessed with an appropriate
measure of caution. There are undoubted successes in certain crop species,
such as tomato (Rick, in press), but the state of genetical knowledge of
these species is vastly greater than that of the groundnut. Genetic maps of
morphological traits in tomato, are well developed, while little progress, if
any, has been achieved in this area for the groundnut. There has been
some progress in producing RFLP maps in Arachis until these can be
related to important morphological (and other) characters they are likely
to remain of purely theoretical interest. Another problem, which does now
seem to be approaching a solution, is that legumes as compared with
members of the Solanaceae are not the most responsive to tissue culture
and in vitro manipulation generally. Until comparatively recently it has not
been possible to obtain regeneration from callus tissue cultures. The
achievement of this is a considerable breakthrough. Nevertheless legumes
in general, and groundnuts in particular, tend to provide highly recalcitrant
The role of biotechnology 715
and intractable material when it comes to in vitro manipulation (Nigam,
1992).
The techniques of achieving transformation of groundnuts are still in the
development phase but there are indications that effective procedures will
be devised. Whether transformed plants will behave as expected is an open
question. Many of us are aware of the genetically transformed arthritic pig
of some years ago. Quite clearly the first stage is to develop transformation
techniques that work; the next task is to identify, extract and clone suitable
genes for introduction. Some genes are already available (the Bacillus
thuringiensis factor might have value in protecting the crop against foliar
feeders) but they are few in number and the great bulk of potentially useful
genes will have to come from other sources. The genus Arachis itself is the
repository for most of the genes whose effects would be valuable if they
could be used to transform A. hypogaea - many pest and disease resistance
genes are in this category. Unfortunately the genetic systems of wild
Arachis species are even less well understood than that of the cultigen; the
identification of useful genes and their attraction for cloning are well
beyond our present capability. To achieve the state of knowledge which
would enable this to be done would require an enormous investment and
the question remains as to whether such an investment would be cost-
effective. There is no guarantee that, if any major resistance genes were
incorporated through transformation, the magnitude of the effect would be
comparable in the donor and recipient species. In practice, the most
effective approach would be to exhaust the genetic resources of section
Arachis species before attempting biotechnological manipulations, in any
but an exploratory way, involving species in the higher order gene pools of
Harlan and de Wet (1971).
The idea that the laboratory will be the source of cultivars in the future is
one which has become lodged in the minds of many, including some of
those involved in making decisions about setting research priorities and
allocating funds. To the politicians, biotechnology has the great attraction
of offering the quick fix. Their time frames seem to coincide with intervals
between elections and there are those in the scientific community who are
prepared to exploit such myopia. However, the short -term advantage is
likely to be counter-productive when eventually the time comes for the
return on investment to be measured and evaluated. The reaction is likely
to be the discrediting of biotechnology as an instrument of crop improve-
ment. A grave error is being made in the implicit promotion of biotech-
nology cum genetic engineering as a substitute for plant breeding rather
than as a potentially valuable supplement to it.
It is becoming increasingly difficult to maintain a spirit of scientific
objectivity and balance in the present age. Although academic freedom
could be regarded by some as a meal ticket for life and justification for a
long-term ego trip exploring esoteric and arcane avenues of research, it did
mean that scientific exploration in the medium and long term could take
716 The future of the ground nut crop
place. Nowadays the short term rules so that results are expected within
three years or so, which can lead to sleight of hand and slick footwork in
what is becoming the sordid business of producing grant applications.
There is, for example, the project-in-hand scenario, where a smooth and
credible proposal can be submitted because the work is already virtually
complete. This type of proposal fares better before a granting committee
than one which seeks to explore difficult areas where problems are ill-
defined and solutions uncertain. The honest and straightforward scientist is
likely to be ever more frequently sidelined by the opportunistic (entrepre-
neurial) operator who may have very little of real value to offer. The field
is open to the plausible charlatan with a superabundant vocabulary of buzz
words who manages to keep ahead of the field and avoids ever being called
to account.
17.5 AGRICULTURAL CONSTRAINTS, RESEARCH AND THE

FUTURE
The International Groundnut Workshop at ICRISAT in late 1991 was an

extremely important occasion for groundnut research workers. The publi-
cation of its proceedings in the following year enabled those who were not
fortunate enough to attend to enjoy the fruits of the presentations and
discussion to the full. In particular, the workshop recommendations
(Nigam, 1992) provide a succinct resume of the general tenor of the
workshop's deliberations. Those of the greatest relevance to the content of
this chapter relate to biotic constraints, current agronomic problems,
genetic resources and germplasm enhancement and utilization.
17.5.1 Biotic constraints

The identification of constraints is a prime prerequisite for implementation
of integrated pest management, the ideals and objectives of which are now
better formulated than in the days 'of the pioneer adaptive researchers in
the 1950s and 1960s. Emphasis was placed on broadening the deepening
biological understanding of the pests and their epidemiology and on close
monitoring of the effectiveness of such management programmes in the
field and the possible development of forecasting systems. The use of
modelling systems in deciding the relative effectiveness of genetic and
management solutions in relief of biotic constraints was advocated. In
practice these would most often be viewed as complementary to each other
rather than as alternatives. The production and distribution of early-
generation hybrids of short-season materials involving rosette-resistant
parentage was recommended, to be distributed over a wide area for
selection to be carried out. This could result in a range of genotypes
carrying genetic resistance with a range of environmental adaptation.
Agricultural constraints, research and the future 717
17.5.2 Current agronomic problems and objectives
Drought resistance and water use economy in the crop are important
aspects (see also Chapter 9) in the context of desertification and increased
frequencies' of drought in the African Sahel. A partial solution to the
problem is to change to shorter season varieties. This could be reinforced
by selection for selectable traits of the root system (which could improve
water uptake) and characters of the shoot system (thicker cuticles, for
example, which could reduce water loss). In addition, physiological
drought resistance and ability to recover from desiccation would be
desirable features to incorporate. These questions are closely related to
high-temperature tolerance and could perhaps form a cluster of selection
objectives. In some areas low-temperature tolerance could be important,
especially in the seedling stage.
Partitioning patterns of assimilates could be manipulated to maximize
seed production, for example, and equally to provide bulk of haulm if this
was desired. With the development of mixed cropping and agroforestry
systems, shade tolerance could become of increasing interest. Soil con-
straints relating to both nutritional deficiencies and acid soil problems
could be the subject of potentially useful study, such as the search for
rhizobial bacteria tolerant to acid soil conditions. The iron deficiency
problem might possibly be relieved by a selection programme on a similar
basis to that suggested for rosette resistance selection.
The plastic mulch technology developed in the Far East (China and
Japan) could be investigated for use elsewhere. It has the advantage in
temperate climates of speeding up soil warming after spring sowing and
more generally of promoting water economy and suppressing weeds. The
plastic residue generates disposal problems but these could be eased, if not
solved, by use of thin films of biodegradable plastics which disintegrate
readily when the crop is mature. The role of the groundnut as an important
component of sustainable farming systems through the incorporation of the
resistance and tolerances mentioned should be explored. The necessity for
appropriate technical training for carrying out these recommendations was
stressed.
17.5.3 Genetic resources and germplasm enhancement

The future of the groundnut crop depends directly on the nature and extent
of genetic resources available for its improvement and adaption to future
contingencies, only some of which can be foreseen. These include adap-
tation to novel systems of production which could evolve in new areas of
cultivation. There can be little doubt that there are extensive areas in the
Americas, for example, where groundnuts could be grown but are not. The
same is true of North Africa and southern Europe: should circumstances
change and it became more attractive to grow the crop under irrigation,
there is little doubt that production could be enormously increased in the
Mediterranean basin. The guided evolution and development of the tradi-
tional farming systems of the developing world could make further
demands on the breeder initially, and ultimately on the pool of genetic
resources.
Resistance to pests and diseases is, as ever, a vital area for further
breeding effort. The expansion of selection at the local level of early
generation hybrid progenies is an interesting and key area in bringing
together the work of breeders at the international agricultural research
institutes and those engaged in adaptive research at the grass-roots level.
This could greatly increase the efficiency of breeding operations in select-
ing for a considerably enhanced range of environmental adaptation and
resistance to or tolerance of local biotic and abiotic problem~ and hazards.
Such selection programmes would not require high levels of sophisticated
training in staff but would rely upon intelligent people endowed with good
commonsense and appropriately trained.
Genetic resources need to be identified which could serve to relieve
problems relating to acid soils and those deficient in micronutrients.
Problems of drought resistance and tolerance breeding could probably be
eased by appropriate use of available germplasm resources, as could
questions relating to high and low temperature tolerances and aflatoxin
contamination. The extent and accessibility of the available genetic
resources in the genus Arachis are considered to be such that biotechno-
logical solutions are the last resort. Arachis hypogaea is really quite
exceptional among legumes in that barriers to interspecific gene exchange
are not as well developed as in many other legume genera, such as Vigna,
at least within section Arachis. Outside this section, but still within the
genus, biotechnology would probably have to be invoked in making inter-
sectional gene transfer. In the current absence of any practically useful
genetic maps, such manipUlations are definitely for the future and not of
immediate relevance.
There is a very real problem to be faced in the dissemination of new and
improved materials and technology. This would clearly be facilitated by
development of a cadre of adaptive researchers acting as an interface
between extension services and the research services of national and
international agricultural research institutes. The transfer of information as
well as materials could be affected by such a chain of communication.
17.5.4 Utilization
Those at the ICRISAT workshop were well aware of the changing patterns
of ground nut utilization. Clearly trends need to be identified and if possible
anticipated, so that appropriate materials can be fed into the farming
systems to maintain their economic viability. This will necessitate market
monitoring and research combined with development of new markets in
References 719
order to maintain demand for the crop and keep the producers in business.
With the present slackening demand for groundnut oil, maintenance of
current levels of production will involve development or expansion of new
and existing outlets - markets with marked potential for adding value to
the basic product. The marketing and processing needs to be arranged so
that the countries of production (and especially the producers) derive a
fairer share of this added value than has been the case in the past.
17.6 A NEW PHILOSOPHY
Research developments to the ground nut since the establishment of the

International Crops Research Institute for the Semi-Arid Tropics have
been particularly exciting to those with prior involvement in groundnut
research in both the developing and developed worlds. At the time of
ICRISAT's inception, the concepts and ideas of the 'Green Revolution'
were very much to the fore. However, 20 years later our approach is much
more broadly based. We have perhaps embraced a philosophy and policy
of 'Green Evolution' based on stable traditional farming systems rather
than the numerous imponderables of tbe Green Revolution and its
aftermath.
Fortunately groundnut research has definable goals and progress
towards them can be monitored and measured. There are many able and
gifted researchers operating in the field and it is to be hoped that the
ultimate objectives of groundnut research, in helping the world's farming
communities which produce the crops, can be reached: to achieve, main-
tain and where possible, improve the level of prosperity and well-being of
the farmers as well as that of the communities they serve.
REFERENCES
Archer, T.C. (1853) Popular Economic Botany, Reeve, London.

Gregory, W.e., Smith, B.W. and Yarbrough, J.A. (1951) Morphology, genetics and breed-
ing, in The Peanut - the Unpredictable Legume, National Fertilizer Association,
Washington DC, pp. 28-88.
Harlan, J.R. and de Wet J.M.J. (1971) Toward a rational classification of cultivated plants.
Taxon, 20, 509-517.
Hemingway, J.S. (1957) The resistance of groundnuts to Cercospora leaf-spots. Empire
Journal of Experimental Agriculture, 25, 60--68.
Hildebrand, G.L. and Smartt, J. (1980) The utilization of Bolivian ground nut (Arachis
hypogaea L.) germplasm in Central Africa. Zimbabwe Journal of Agricultural Research,
18,39-48.
Johnson, F.R. (1964) The Peanut Story, Johnson Publishing Co., Murfreesboro, North
Carolina.
Klozova, E., Svachulova, Smartt, J. et al. (1983a) The comparison of seed protein patterns
within the genus Arachis by polyacrylamide gel electrophoresis. Biologia Plantarum, 25,
266-272.
Klozova, E., Turkova, V., Smartt, J. et al. (1983b) Immunochemical characterization of seed
proteins of some species of the genus Arachis L. Biologia Plantarum, 25, 201-208.
Kochert, G., Halward, T., Branch, W.D. and Simpson, C.E. (1991) RFLP variability in
peanut (Arachis hypogaea L.) cultivars and wild species. Theoretical and Applied
Genetics, 81, 565-570.
Krapovicakas, A. (1969) The origin, variability and spread of the groundnut (Arachis
hypogaea), in The Domestication and Evolution of Plants and Animals, (eds P.J. Ucko and
G.W. Dimbleby), Duckworth, London, pp. 427-441.
Nigam, S.N. (ed.) (1992) Groundnut - a global perspective. Proceedings of an International
Workshop 25-29 November 1991 ICRISAT Centre. ICRISAT, Pantancheru, Andra
Resslar, P.M. (1980) A review of the nomenclature of the genus Arachis L. Euphytica, 29,
813-817.
Rick (in press) in Smartt, J. and Simmonds, N.W. (eds) Evolution of Crop Plants (2nd edn)
Longman, London.-
Singh, A.K. (1988) Putative genome donors of Arachis hypogaea (Fabaceae), evidence from
crosses with synthetic amphidiploids. Plant Systematics and Evolution, 160, 143-151.
Smartt, J. (1978) Makulu Red - a 'Green Revolution' groundnut variety. Euphytica, 27,
605--608.
Smartt, J., Gregory, W.C. and Gregory, M.P. (1978) The genomes of Arachis hypogaea L. 1.
Cytogenetic studies and putative gene donors, Euphytica, 27, 665--675.
Stalker, H.T. (1991) A new species in section Arachis of peanuts with a D genome. American
Journal of Botany, 78, 630--637.
Stalker, H.T. (1992) Utilizing Arachis germplasm resources in Groundnut - a Global
Perspective, (ed. S.N. Nigam), ICRISAT Patancheru, A.P., India, 281-295.
York, E.T. and Colwell, W.E. (1951) Soil properties, fertilization and maintenance of soil
fertility, in The Peanut - the Unpredictable Legume. National Fertilizer Association,
Washington DC, pp. 122-172.
Index
a-conarachin 183 screening 530

Abiotic stress 128 teratogenicity 509
Absorption, gut 183 toxicity 509,516,520,522-3,525-8
Acacia 258 Aflatoxins
Acarina, mites 414 Bl (AFB l ) 510-11, 514-29, 532-4, 538,
Acetylene reduction 268-9 540
ACIAR605 B2 (AEB 2) 510, 515, 517-18, 532-3, 538
Adaptation (rhizoma peanuts) 637-8 G l (AFG l ) 510, 515, 517-18, 532, 534
Adaptive research 675-98, 705, 706 G z (AFGz) 510, 515, 517-18, 532-3
in Zambia 675-95 Ml (AFM l ) 516-17, 534, 540
Additive effects, insect resistance 571, 574 Africa 3,4,6,449,575-85
Additives, peanut butter 490-1 Agricultural constraints 716-19
Adenosine Agroforestry systems 667
diphosphate (ADP) 255 Agronomic practices 680-90
monophosphate (AMP) 266 Agrotis spp. 407
triphosphatase (ATP-ase) 267 Alachlor 642
triphosphate (ATP) 255, 256, 266 Alanine 186, 191
Aeschynomene 260, 653 Aldicarb 374
Aeschynomene indica 374 Aldrin 454
Aflatoxicosis Aleyrodidae, whiteflies 411
animals 522-4 Allergens 184
humans 524-8 Allergy 184
and hepatic cancer (primary liver Allozymes 575
cancer PLC) 527-8 Alternaria Jeafspot (Alternaria spp.) 357-85
hepatic cirrhosis 526-7 Alternate branching 63-4
kwashiorkor 525-6 Aluminium (AI) 202
Reye's syndrome 527 Ambinervosae (section) 98, 100, 108, 112,
Aflatoxin 360-2, 455, 509 et seq., 568-749, 119
615,631,704 Amendoim 31
analytical methods 528-35 Amerindian vernacular names 31
carcinogenicity 509,516-18,524,527-8 Amino acid composition, groundnut protein
chemistry 517-21 185
contamination control 535-9 Amsacta spp. 413
contamination, post-harvest 513-14 Anarsia ephippias 414
contamination, pre-harvest 511-13 Anatomy 75-88
detection 528-35 embryo development 84-8
destruction 537-9 leaf 78-80
metabolism 517-21 reproductive bodies 80-4
mutagenicity 501 root 76
natural occurrence 510-11 seedcoat 86
prevention 535-6 stem 76-8
quantitative measurement 530-5 wild species 86-8
removal 536-7 Anchic 31
sampling 529-30
722 Index
Ancistro termes latinatus 400 A. hoehnei 442
Anthracnose, see Colletotrichum spp. A. hypogaea 32-3, 45, 53 et seq., 100-2,
Antibiosis 432 106, 110,112-13, 122, 442, 562-3,
Anti-nutritional factors 190, 203 627-35,695
Antixenosis 432 see also Groundnut
Ants 445, 471 subsp. fastigiata 55,553,692
Apanteles 447 var. fastigiata 55, 553-4
Aphanus sordidus 414 var. vulgaris 55, 553-4
Aphasmatylenchus 372 subsp. hypogaea 54,553,692
Aphasmatylenehus straturatus 376-7 var. hirsuta 55, 553-4
Aphelenehoides 372 var. hypogaea 54, 55, 553-4
Aphelenehoides arachidis 377-8 A. ipaensis 604
Aphididae, aphids 410,571,586,591,598, A. kemf-mercadoi 442
601 A. khulaminii 442
Aphis eraecivora 410 A. lignosa 44
Aphis glycines 410 A. luteseens 45,51-2, 100, 119
Aphis gossypii 410, 566 A. macedoi 44
Aphis robiiae 410 A. marginata 45,50, 100, 119
Aphis solanella 410 A. martU 45, 48-9, 100, 122
Apparent root depth progression (ARDP) A. monticola 45,47,100-2,105-6,
287-8 109-10, 112-13, 122, 126
Application of fertilizer 229,230,233, A. nambyquarae 45,98
235-9,241,243 A. oteroi 44
Applied ecology 415 A. paraguaniensis 45,48, 100, 128,442,
Aproaerema modicella, groundnut leaf 560
miner 413,417,419-21,427, A. pintoi 44, 645-55
447-9,454 A. prostrata 45,50, 100, 119
Arachidins I, II and III 513 A. pseudovillosa 442
Arachidonic acid 178, 181 A. pusilla 45, 53, 98, 100, 126,442
Arachin 183-4 A. repens 45, 49, 50, 100, 126,560
A rachis (genus) 31-2, 625 A. rigonii 45, 49, 100, 128,442
see also sections and species A. spegazzinii 102, 106, 109, 113
Arachis (section) 98,100,102-5,107,109, A. spinaclava, see A. glandulifera
113, 119, 122 . A. stenocarpa (A. stenosperma) 44, 104,
Arachis species 126, 560, 564
A. afrieana 54 A. stenophylla 442
A. angustifolia 45, 100 A. sylvestris 119
A. appressipila 442,560 A. tuberosa 45,47,95,100,108,119
A. asiatica 55 A. valida 442
A. batizoeoi 45,46, 100-4, 106, 109, 113, A. villosa 45,46,98, 100-2, 122,442
442,560 A. villosa var. eorrentina, see
A. X batizogaea 45, 98, 100,442 A. eorrentina
A. benthamii 45,48, 100 A. villosulicarpa 45,51, 100, 122, 126,
A. burkartii 45, 50, 100 128,560
A. cardenasii 102, 125, 126,342,442, Arb (cv) 636, 640, 643-4
560,572 Arbrook (cv) 636,641-2,644-5
A. chaeoensis (A. chacoense) 98, 126, Arblick (cv) 640, 643-4
442, 560, 564 Archaeological evidence 115
A. ehiquitana 442 Argentina 2,4,6,613-14
A. correntina 45,98, 100-3, 125, 442 Ash content 201
A. diogoi 45,46, 100, 119 Ash grey cotton weevil (Myllocerus
A. duranensis 102, 104, 106, 109, 113, undecimpustulatus maeulosus) 406
342, 442, 560, 572 Asia 3,5,6,446-9
A. glabrata 45,52,98, 100, 102, 126,442, Asparagine 185, 269
560,633,636-7,654-5 Aspartic acid 185-7, 190
A. glandulifera 100, 102 Aspergillus 568, 572, 599
A. guaranitiea 45,47,48,100,108 resistance to colonization 568-9,572-3,
A. hagenbeekii 45,52-3, 100, 126,442, 607
560,633 resistance to toxin production 568-9, 612
A. helodes 45-7, 100 Aspergillusfiavus, yellow mould 360-2,400,
723
414,510-13,517,568-73,591,598, Bracon 447
601,612,614,684 Bradyrhizobium 260 et seq., 624, 638, 642,
in vitro seed colonization 569, 572 650,651
resistance in wild species 572 Branching pattern 63-4
toxin production on defined media 572 Brazil 2, 4, 6, 613, 614
Aspergillus niger, crown rot 454,511,693 Breeding methods 574-5, 577, 586
Aspergillus parasiticus 360-2, 510, 517, Breeding objectives 577,586, 591, 598, 605,
568-9, 570-1, 684 605
Atherosclerosis 181-2 Brunswick grass 645
Attagenus fasciculatus, black carpet beetle Bud necrosis virus (BNV) 565, 566, 591,
461 598
Australasia 606 Buprestidae, jewel beetles 402
Australia 606 Burkina Faso 3, 4, 6, 576, 580
Availability of amino acids in products 190 Burundi 5, 6
A vena sativa, oats 640 B-vitamins 200
Avga choaspes 447
Avoidance of pests 432-3,459--60 Cacahuate, cacahuete 31
Axonopus affinis 643, 652 Cadra cantella 462
Calcium (Ca) 202, 214, 215-26, 230-3
[3-carotene 200 deficiency correction 223--6
[3-oxidation pathway - fatty acids 266 deficiency symptoms 220-1
Bacillus thuringiensis 464 needs 156, 215, 221-3
Backcrossing 574 needs in relation to genotype 217
Bacterial needs for seed development 216 et seq.
diseases 336-9 sources 223--6
leafspot (Xanthomonas spp.) 338-9 dolomite 226
wilt (Pseudomonas solanacearum) 336-8, gypsum 223-4
693 limestone 223-5
Bacteroids 262,266,267 phosphogypsum 223
Bangladesh 591, 598 status 296-8
Beak (pod) 74 Calixin 352'
Belonolaimus 372, 375 Cambodia 601,605
Belonolaimus longicaudatus 375--6 Cameroon 3, 4, 6
Bemisia tabaci 411 Canopy development 302, 305
Benefin 642 Captafol 371
Benin 3, 4, 6 Carbaryl 453
Benomyl 352, 354, 356, 364 Carbofuron 374, 567
Bentazon 642 Carbohydrate content 196-8, 205
Bhutan 591 Carbosulphan 454
Bicarbonate 273 Carboxin 360
Biochemistry, seed 149-53 Carcinogenicity of aflatoxin 509,516-18,
Biomass, total (W) 301 520-1, 524, 527-8
Biotechnology 612,618,714-16 Cardiophorus subspinosa 406
Biotic constraints 716 Cardiophorus cognatus 406
'Biteability' 441 Carotenoids 200
B1ackhull (Thielaviopsis basicola) 363-4 Caryedon serratus, groundnut pruchid 415,
Blue couch grass 643 461-2
Bolivia 613, 614 Carpophore, see Peg
Bordeaux mixture 352, 355 Casein 193-4
Boron (B) 201, 234-8 Cassia fasciculata 374
deficiency 234-8 Caterpillars 692
toxicity 238 Caulorhizae (section) 98, 100, 103, 107-8,
Botanical types of Arachis hypogaea 353 112, 119,645--6
Botrytis blight (Botrytis cinerea) 364 CBR 227
Brachiaria 649 cDNA 105--6
Brachiaria brizantha 652 Cellulose 199
Brachiaria decumbens 652 Celosia argentea 455
Brachiaria dictyonema 652 Central Africa 5,6
Brachiaria humidicola 652 Centres of diversity of cultigen 113, 115-18
Brachiaria ruziziensis 652 wild taxa 113-18
724 Index
Centres of origin of Arachis taxa 112-17 Cotton (Gossypium hirsutum) 559
Centrosema 624 Cowpea strains (Bradyrhizobium) 256, 263
Centrosema macrocarpa 650 Creontiades 412
Cercospora arachidicola, early leaf spot Creontiades pallidifera 412
346-8,557,558,586,591-8,606-7, Creontiades pallidus 412
644, 653, 690, 692 Crickets 408
Cercosporidium personatum, late leaf spot, Criconomella372, 375
see Phaeosariopsis personatum Criconomella ornata 375, 376
Chalimbana (cultivar) 676-8, 691 Cristulariella maricola, zonate leafspot 358
Charcoal rot (Macrophomina phaseolina) Crop growth rate (CGR) 301
364-5 Crop phenology 319
Chelonus 447 Cropping systems 673-5
Chemotaxonomy 105--6 Crude fibre 199
cDNA 105--6 Cultivar development 612
flavonoids 105 Cultivar transpiration efficiency 309-10
isozymes 105 Cultivars released in
RFLP 106 Africa 580-5
seed proteins 105 East Asia 579,586-8,588
Chickpea flour 195 India 592-5
Chile 614 southern Asia 588-91, 600-1
China 3,5,6,446-9, 579-87, 588-9, 601 Cultivation in new geographic areas 706
Chitemene system 667 Cultivation systems 666-75
Chlorine (CI) 202, 238 agroforestry 667
Chloris gayana, rhodes grass 643 ley farming 667, 679-71
Chlorothalonil 350, 352, 354, 364, 371, 446 permanent cultivation, irrigated 667,672-3
Chlorpyrifos 454 permanent cultivation, rainfed 667, 671
Choanephora spp., choanephora leaf spot semi permanent cultivation 667, 668-9
·358-9 shifting cultivation 667-8
Cholesterol 182 Cultural control (pests) 454--61
Chromium (Cr) 201, 202 Curculionidae, weevils 406
Chromosome complements 104 Cylindrocladium blackrot (CBR)
Chrysomelidae 406 (Cylindrociadium crotalariae) 227,
Cicadellidae, jassids 411, 427 368-9,568,606-7,612,653
Climatic change 704-5 Cystine 183, 186, 190, 194
CNAR576 Cytotaxonomy 101
Cobalt (Co) 202
Coccinellids 449 Damage by foliar pathogens 558
Cold tolerance 599 Damage estimation of foliar feeders 424-5
Coleoptera 402, 450 Decortication 486
Collecting expeditions, germplasm 118-22 Defatted flour 188
Collections, germplasm 118 et seq., 586 Deficiency, minerals 220-43
Colletotrichum arachidis 365 B 234-8
Colletotrichum dematium (truncatum) 365, Ca 220-5
653 Cu 238
Colletotrichum gloeosporioides 653 Fe 233,239
Colletotrichum mangenoti 365 Mn240
Colombia 2, 4, 6 P 228
Companion species (forage Arachis) 652 Zn243
Composition, kernel 173-4 Deficit, water 284-7,305
Composition, oil 176-9 Defoliation damage simulation 425-6
Conarachin 183 Dermaptera, earwigs 400, 401
Conarachin I 184 Descriptors 123, 124-8
Conarachin II 184 Desertification 577,586,591,605,606,614
Conservation of germplasm 129 Desmodium 624
Constriction, pod 74-5 Destruction of aflatoxin 537-9
Copper (Cu) 201, 202, 238-9 Detection of aflatoxin 528-35
Corcyra cephalonica 462 Detection of resistance to foliar pathogens
Corn earworm (CEW) 439, 445 559--60
Cornflour 194 Deterioration of seed 162-5
Costelytra zealandica 440 Development
725
of canopy 302, 305 East Asia 3, 5, 6, 446-9, 579--87, 588-90,
of pod 72-3, 140-4 601
of seed 73-4, 144-9,216-25 Ecology 114
Diabrotica undecimpunctata howardi, applied 415
spotted cucumber beetle, southern Economics of ecological control 415, 422
corn rootworm 406,421,453,463, Ecuador2,4,6,614
468,571 Egypt 5,6
Diacrisia obliqua 413 Eicosanoic acid 178
Diaecodorus 406 Elasmolomus sordidus 414
Diaminobenzidine (DAB) 268 Elasmopalpus 453
Dibromochloropropane (DBep) 377 Elasmopalpus lignosellus, lesser cornstalk
Dichlorvos 427, 453 borer 571
Dicloran 364 Elateridae, click beetles, wireworms 406,
Dietary fibre 199 407
Diet supplementation 194 Eleusine coracalla, finger millet 674
Diffusion barrier, nodules 267 Empoascafabae 411,644
Difiubenzuron 427 Empoasca kerri 410,411,626
Digestibility 191 Encyrtidae 401
Digitaria 460 Endocytosis 262
Digitaria didactyla, blue couch grass 643 Endosulphan 453
Digitaria eriantha, angola grass 643, 652 Enlleothrips flavens 410
Dimethoate 427, 430, 447, 452, 453 Environmental effects on yield 308
Diplodia gossypilla, diplodia collar rot Environmental factors affecting
365---{i pest populations 420-2
Disaccharides 196, 197 root growth 289-93
Disease reduction 632-3 Environmental stress effects
Disease resistance 126, 128, 560-1, 564 on nitrogen fixation 273-5
Diseases 336-78 on nodulation 272
bacterial 336-9 Enzyme-linked immunosorbent assay
fungal 346-72 (ELISA) 529, 532-5
nematode 372-8 Erectoides (section) 98, 100, 104, 107, 108,
viral 339-46 112, 119, 654---{i
Dithane 694 Essential amino acid deficiences 191
Ditylellchus 372 Essential amino acids in products 189
Ditylellchus destructor 377-8 Establishment of forage species 650-1
Diversity in products 700-2 Ethnography 35---{i
DNA 105---{i Etiella behrii, lucerne web moth 407
Dormancy, seed 153-5 Euborella allllulipes 400
Dorylus labitus 402 Euborella plebeja 400
Dorylus orienta lis 402 Eucelatoria armigera 445
Drought 577, 586, 591, 601, 605, 606, 614 Eurhizomatosae (series) 633, 636, 637,
effects of 309 639-41, 643-4
intercropping 322-3 Europe 3, 4, 5
management 317-4 Evolutionary history (Arachis) 710-11
mulching 323-4 Exorista xanthopis 447
plant population density 319-22 Expression of oil 497-505
supplementary irrigation 317-18 Extraction methods for oil 497-505
tolerance 310-17, 319, 578
Drying, crop 484-5 Fabaceae, see Leguminosae
Dry matter, pod 307 False wireworms (Tenebrionidae) 406, 407,
Dry matter, total (TDM) 306-8 468
Dyanap 696 Fallowing systems 667-9
Dysmicoccus brevipes 401 FAO 576
Farming groundnuts 702-9
Early leafspot (Cercospora arachidicola) Farming systems 664
346-8, 558, 577, 586, 591, 598, Fast growing nodule bacteria (Rhizobium)
606,612 260
Early maturity 605 Fastigiata (subsp.) 55, 99, 123, 553, 556,
Earwigs (Dermaptera) 400, 401, 445 557,692
East Africa 56 Fastigiata (var.) 55, 553, 554
726 Index
Fat absorption 183 enhancement 717-18
Fatty acids 174, 176-7 utilization 131
FAW 576 Ghana 3,4,6
Feltia 407 Globulins, seed 183
Fermented flour 193 Glucose 185
Fermentation (fungal) 189-92, 198-200 Glutamic acid 185-7, 190
Fertilization (floral) 70, 138 Glutamine 185
Fertilizer use 707 Glutamine-2-oxyglutarate amino transferase
Fibre, dietary 199-200 (GS:GOGAT system) 263-4
Filtration (oil) 502-5 Glycine 186, 191
Fixation, nitrogen 227, 255 et seq., 287 Glycine max, soybean 559
Fixation, phosphorus 228, 229 Glycolipids 178
Flavonoids 105 Goitrogens 190, 205
Flavour (products) 487, 490, 491, 493, 494 Goniozus 447
Flight activity (insects) 415-16 Gossypium hirsutum 559
Floral morphology 66-9 Graphognathus leucoloma, white fringed
Flower colour 125 weevil 406
Flowering 66,69-70, 138,213 Grasshoppers 408
Flowers 66 Grey cotton weevil (Myllocerus
Fluazifobutyl 642 undecimpustulatus maculosus) 406
Forage feeding value 640, 649-51 Groundnut
Forcipula quadrispmosa 400 anatomy 75-89
Formicidae, ants 401,445,471,571 biosystematics 96-109
Frankliniella fusca 409, 644 breeding 552-616
Frankliniella occidentalis 409 chlorosis (Aphasmatylenchus straturatus)
Frankliniella schultzei 409, 410 376-77
Free amino acids 197 composition 173-205
Free fatty acids 179 development biology 138-67
Free sterols 178, 179 diseases 336-78
Fructose 185 farming systems 664-99
Fruit (pod) 72-3, 140-4 forages 624-57
Functional nodules 263-8 future 700-19
Fungal diseases of stem, root and pods genetic resources 109-33
359-72 history 26-39
Fungal foliar diseases 346-59,558-65 improvement 709-13
Fungicides 348,350, 352, 354-6, 360, 363, leaf miner (Aprogerema modicella) 413,
365 427, 439-41, 468
Fusarium solani 346-71 meal 188
mineral nutrition 214-5
-y-methylene glutamic acid 185 morphology 43-75
-y-methylene glutamine 185 mycotoxins 509-40
Galactic binding (GL) 264 nitrogen fixation 255-76
Gambia 3, 4, 6 nutritive value 173-205
Gene pools 110-12 oil 480-6
Genetic origins 26-39
factors in root growth 288-9 pests 395-469
maps 575 processing 480-508
resistance 711-12 production 1-22
resources 109-10, 717-18 reproductive biology 138-67
Genome analysis 103 rosette virus 344,565,577,692-3
Genomes A, B, D 98, 101, 103 streak necrosis (sunflower yellow blotch
Genotype improvement in drought virus) 410-11
tolerance 310-12,318-19 trade 1-22
Geographical utilization 480-508
dispersal/distribution 105, 108-9, 112-13 water relations 281-325
origins, wild taxa 636, 647 see also Arachis hypogaea
Germplasm Growth habit 125
collecting 554, 607, 615 GTZ576
collections 554-5 Gums 199
conservation 129 Gypsum 223-4
727
Habitats, wild species 114 Hypogaea (var.) 54, 55, 99, 123,553,554
Haemagglutinins (lectins) 190,205,263,264
Hairy caterpillars 468, 591 Ibimani 31
Hairiness Ichneumon 447
leaflet 66 ICRISAT 576,605,607
peg 71 Immunochemical methods (aflatoxin
stem 61 detection and measurement) 532-5
Harvest index (H) 301 Immunosuppression 524
Harvesting 483 Inchic 31
Haulm India 3, 5, 6, 446-9, 591, 593-5, 598, 601
feeding value 632 Indian Council of Agricultural Research
quality 627-9 (I CAR) 599
use 625---{5 Indian meal moth (Plodia interpunctella)
yield 629-30 462
Hay production 630-1 Indigofera hirsuta 374
Hay storage 631 Indonesia 601,605
Health implications, oils 181 Induced mutagenesis 601
Helicoverpa 451, 571, 574, 586, 591, 601 Ineffective nodules 260,265
Helicoverpa armigera 413,426,451,464 Infection pathways 256,262
Helicoverpa punctigera 413 Inflorescence 66
Helicoverpa zea 413, 416, 425 Inoculum, rhizobial 226
Heliothis armigera 452 Insecticides 427, 450-4
Heliothis virescens, tobacco budworm 416, Insoluble carbohydrates 196-8
445 Integrated Pest Management (IPM) 397,
Heliothis zea, corn earworm (CEW) 441, 398,463-7
445,463,468,644 Intercropping 454-5
Helopeltis 412 International Rand D support 576-7
Hemicellulose 199 Intersectional hybrids 104
Hepatic cancer 527-8 Interspecific hybridization 103 et seq.
Hepatic cirrhosis 526-7 In vivo aflatoxin detection 529, 532-5
Herbicides, pre-emergence 642,694 Ipomaea Jistulosa 455-7
Herbicides, post-emergence 642 IRAT 576
Heteronyx arator 403, 440 IRHO 576, 580, 582, 583
Heteronyx piceus 403 Iron (Fe) 202, 233, 239-40, 273-4
Heteropogon contortus, native black Irrigation management 458-9
speargrass 643 Isoleucine 185, 186, 188, 193
High performance liquid chromatography Isotopes 270
(HPLC) 532, 535 Isozymes 105
High temperature tolerance 586 ISRA 576
Hilda patruelis 401,449,468,573,577 Italian ryegrass (Lolium multifiorum) 640
Hirsuta (var.) 55, 99,123,553,554 Ivory Coast 3, 4, 6
Histidine 185, 186 IVSCAF, in vitro seed colonization by
Historical industrial development 36-8 Aspergillus fiavus 569, 572
History, groundnut 24-42
Hodotermes mossambicus 400 lacobiasca formosana 411
Hogging off 625 Japan 3, 5, 6, 446-9, 579, 587, 589
Hollow heart 235 J assids (J assidae/Cicadellidae) 411, 427,
Holotrichia serrata 406, 421 468,571,574,577,591,598
Host plant resistance 430-41
Hybridization 574 Korea(s) 3, 5, 6, 446-9, 579, 590, 591
Hydraulic pressing 498-500 Kwashiorkor 195, 525---{5
Hydrocarbons 178, 179
Hydrogenation 501-2 Labidura riparia 445
Hymenoptera 401,445 Laboratory feeding experiments (insects)
Hypercholesterolaemia 182 422
Hyperparasites 446-50 Lacewings 449
Hypochthonellidae 401 Land management systems 665
Hypochthonella caeca 401 Laos 601
Hypocotyl 59 Larval damage estimation 424-5
Hypogaea (subsp.) 54, 99, 123,553,554 Lateral roots 59
728 Index
Leaf Manobi 31
area development 285 Mantids 408
display 286 Maturity indices, seed 153
hoppers, see Jassids Meal, groundnut 21
miner (Aproaerema modicella) 413, 417, Mealy bugs (Pseudococcidae) 401
419-21,427,441,447-9,454,571, Measurement of nitrogen fixation 268--72
574,577,591,598,601 Mechanisms of seed deterioration 163-5
morphology 64--6, 125 Medicago 624
relative water content (RWC) 282 Medicago sativa 640, 641
senescence 286-7 Megagametophyte 82
spots (cercospora) 346-50, 35~1, 564, Megasporogenesis 82
577-8,586,591,598,601,605, M egalurothrips usitatus 408
606-7 Mehlich 1 extraction 222, 228, 230--2
water potential 283 Mehlich 3 extraction 231
Leaflet morphology 64, 65 Melanosis (Stemphylium botryosum) 358
Lectins (haemagglutinins) 190,205,263, Meloidogyne 372-5, 645
264 Meloidogyne arenaria 373-5, 586, 645, 654
Leguminosae 97, 255 Meloidogyne hapla 373-5,586,645,654
Lepidiota 429 Meloidogyne incognita 373-5, 645, 653
Lepidoptera 407,413,571 Meloidogyne javanica 373-5,645
Leptosphaerulina arachidicola (pepper leaf Mesoleurus dentipes 406
spot) 653 Metabolism, seed 152-3
Leptosphaerulina brislanna 645 Meta-sodium 372
Leptosphaerulina crassiasca 354-5 Metasystax 453
Leucine 186 Methionine 183-8, 190--4
Ley farming 667-9 Mexico 2, 4, 6
Life form 126 Micronutrients 234-64
Lignin 199 Microsporogenesis 80--2
Linoleic acid 177 Microtermes 400, 421
Linolenic acid 178 Mineral nutrition 161 et seq.
Upaphis erysimi 410 Miridae 412, 468
Lipid metabolism 264 Mixed cropping 673-5,707-8
Lipopolysaccharides (LPS) 262 Molecular techniques in groundnut
LISA (low input sustainable agriculture) improvement 612
615 Molybdenum (Mo) 202, 242
Lorsban 453 Monkeys 577
Lotus pedunculatus 637 Monocrotophos 452
Lucerne/alfalfa (Medicago sativa) 640, 641 Monosaccharides 196, 197
Lutein 200 Morphology 43 et seq.
Lygaeidae 413 Mott dwarf elephant grass, Pennisetum
Lysine 184-8, 190--5,205 purpureum 643
Mozambique 5, 6
Macrophomina phaseolina, charcoal rot Mucilages 199
364-5 Mulches (synthetic) 457-8
Macroptiliurn 624 Mulching 323-4,455-7
Macrosiphon avenae 410 Multiple pest and disease resistance 371,
Macrosiphon euphorbiae 410 374
Madagascar 5,6 Munduvi 31
Magnesium (Mg) 202, 214, 231-3 Mutagenicity induced by aflatoxin 509
Maize (Zea mays) 559 Mutation affecting oil composition 177
Malawi 5, 6, 576, 581 Mycosphaerella arachidis, see Early leafspot
Malaysia 601,605 Mycosphaerella berkeleyii, see Late leafspot
Male gametophyte 80--2 Myanmar 3,5,6,446-9,591,598,601
Mali 3, 4, 6 Myllocerus undecimpustulatus maculosus
Mancozeb 354-5 ash grey/grey cotton weevil 406
Mandovy 31 Myriapoda, millipedes 407, 408, 468, 571,
Mandubi 31 577
Mandues 31 Myrothecium roridum, groundnut leaf
Manganese (Mn) 202, 240--1 blight 358
Mani 31 Myzus persicae 410,566
729
Nasutitermes 400 Orthoptera 408
Natural abundance 15N method 271-2 Osmotic potential 282
National Peanut Foundation 612 Osmotic pressure 283
Natural occurrence of aftatoxins 510--11 Ox amyl 374
Neem cake mulch 456
Nemagone 567 Pachycondyla 449
Nematicides 368, 374, 376, 377 Pakistan 3, 5, 6, 446-9, 591, 596, 598
Nematodes 372-8, 577, 586 Pangola grass (Digitaria eriantha) 643
Nepal 588, 597, 598 Panicum maximum 652
Net protein utilization (NPU) 191, 194 Papilronoideae 97, 255
Neutral detergent fibre 199 Paraguay 2, 4, 6, 613, 614
Nezara viridula 426 Paraplanobia 414
Niacin 200 Parasitica (Hymenoptera) 445
Niger 3,4,6 Paribaea orbata 446
Nigeria 3, 4, 6, 576 Parkia 258
Nitrogen 214,226-8 Parkia ciappertoniana 460
assimilation in plants 257 Partitioning and harvest index 301-2
fixation 227, 255 et seq., 287, 706 Partitioning assimilates to pods (P) 301
fixation, measurement 268-70 Partitioning rates (PR) 302
reduction 255 Paspalum dilatatum, dallis grass 643
Nodulation in Arachis 257-60 Paspalum nicorae, brunswick grass 643
Nodulation genes 261,262 Paspalum notatum, Pensacola bahiagrass
Nodulation, promiscuous 260 374,641,643,655
Nodule characteristics 256 Peanut
Nodule structure 265-7 butter 188, 480--1
Nodules additives 490--1
effective 260, 263-8 processing 488
functional 263-8 production 487-9
ineffective 260, 265 production equipment 488
rhizobial components 266 candy 494-6
Nomenclature (taxonomic) 97 et seq. clump virus (PCV) 343-4, 565, 567, 577
Non-essential amino acids 186 CRSP614
Non-reducing sugar 197 mottle virus (PMV) 339-40, 565, 566,
North America 2,4,6,441-6,607-13 577,655
Nuclear polyhedrosis virus (NPV) 445-6 stripe virus (PSTV) 340--2, 565, 566, 586,
Nutrition, calcium 156 587,601
Nutrition, mineral 161 stunt virus (PSV) 342-3, 653
Nutritional quality (seed) 128 Pectin 199
Nutritional value/chemical composition 215 Pedigree method 574
Nutritive value 173 et seq. Peg 70--1
Nuvacron 694 entry 298-308
Pegging and podset 293-300
Oats (A vena sativa) 640 Penman equation 308
Odontotermes 414,455 Pennisetum purpureum 461,643
Odontotermes brunneus 400 Pentatomidae 413
Odontotermes obesi 400 Pepper spot and leaf scorch
Odontotermes wallonensis 400 (Leptosphaerulina crassiasca) 354-5
Oidium arachidis (powdery mildew) 356-7 Perennes (series) 98, 100, 107
Oil Peri bacteroid membrane (PBM) 267
and fatty acids 174 Peribacteroid unit 267,268
composition 176-9 Permanent cultivation, irrigation 667, 672-3
content 175, 176,205 Permanent cultivation, natural rainfall 667,
Crops Research Institute, Wuhan, China 671-2
586 Pestalotiopsis leafspot (Pestalotiopsis
expression 497-505 arachidis, P. adjusta and P.
quality 179 versicolor) 356
Oleic acid 177 Pests and diseases (forage species) 653-4
Oleic/linoleic ratio 179 Phaeoisariopsis personata, late leaf spot
Oleosomes 264-6 348-50,558,577,586,588,598,
Origin of cultigen 113, 115, 117 601,605,606,607
730 Index
Phascolus 455 Processing (groundnut) 188,482
Phenamiphos 374 Procumbensae (section) 98,100,111,119,
Phenolic acids 202, 203 122,654-5
Phenylalanine 185--6 Production data 552
Pheromone traps 418 Production technology 666
Philippines 601, 605 Productivity, farming 703
Phoma blight 356 Products 188
Phoma arachidicola 577 Proline 185, 186, 190, 191
Phoma microspora 356 PROMANI612
Phomopsis 653 Promiscuous nodulation 260
Phorate 454 Prorhizomatosae (series) 98, 100, 107, 112,
Phospholipid 178 119,633
Phosphorus (P) 202, 214, 226, 228-30, 234, Protection in storage 462
274,275 Protein
Photosynthesis 284 efficiency ratio (PER) 19J)....6
Phyllophacia congrua 440 isolates 189 •
Phyllosticta (Phyllosticta spp.) 357, 645, 693 polymorphism 183
Phymatotrichum omnivorum 645 quality 191, 192
Physical resistance 432 quality in storage 196
Phytic acid 190, 200 Pseudococcidae 401
Phytoalexins 513 Pseudococcus solani 401
Phytosterols 179 Pseudomonas solanacearum 568, 587, 601
Pistache 31 Pseudoplea 693
Plantations 667 Psyllechthrus oophagus 401,449
Plant Puccinia arachidis, rust 350-3, 558, 577,
analysis 329 586-9, 601, 605-7, 612, 616, 642,
breeder's role 711-13 653
populations 157-8, 319-22, 680, 684-8 Pure line selection 586
water potential 282 Pyralidae 462
Planting rates (forage groundnuts) 641-2 Pythium diseases 366-8
Planting times (forage groundnuts) 642 Pythium myriotylum 373, 568
Platythyrea 449
Pleospora 693 Quality retention 485
Plodia interpunctella, Indian meal moth 462 Quantitative analysis for alfatoxin 530-1
PNUTGRO model 286 Quercetin glycosides 441
Pod 72-5, 125 Quinalphos 453
dry matter 307
filling 300, 301 Rancidity 179-81
filling phase duration (t) 301 Randomized groundnut oil 182
and root knot disease 373-5 Raffinose 196-8
and root lesion disease 375-{i RAPD 175
yields 306-8 Reaction scoring (disease) 559
Pollination 70, 138 Recurrent selection 574
Polymixa graminis 566 Reduced rainfall effects 575
Polyploids 80 Reducing sugars 197
Polyploidy 104 Refining (oil) 501
Post-Columbian history 26-30 Regional progress 575--615
Post-emergence herbicides 642 Relative humidity effects on storage 485--6
Post-harvest pests 414 Relative water content (leaf) RWC 282
Potassium (K) 202, 214, 230-2 Repellance 432
Potential yield losses 155-8 Reproductive
Powdery mildew (Oidium arachidis) 356-7 axis (R) 66
Pre-emergence herbicides 642 biology 138 et seq.
Pre-historic cultivation 25--6 parts 80-4
Pre-soil penetration (fruit) loss 155--6 sinks 301
Press cake 480 Requirements for
Press extraction 188 Ca 215
Prevention, aflatoxin formation 535--6 Mg232
Prices 15-17 P228
Principles of seed storage 162-3 Resistance, diseases, 98, 126, 128, 558-{i4
731
foliar pathogens 98, 126,560,561,564 Saponifiable fraction (oils) 176-8
nematodes 103 Saponins 190, 202
soil-borne pathogens 567-8 Scab (Sphaceloma arachidis) 355-6
toxin production 568-9, 572-3, 612 Scarabeidae, white grubs, cockchafers, May
yellow mould colonization 568-70,572-3, bugs, June beetles, 402, 406, 419,
612 468
Resistance, host plant 430-45 Scenopenidae 450
Resistance, pests Scirtothrips aurantii 408
aphids 433-4, 566, 577 Sclerotinia blight (Sclerotinia minor,
black carpet beetle 462 Sclerotinia sclerotiorum) 370-1,
corn earworm 439, 445 568,606-12
groundnut leaf miner 439-41 Sclerotium rolfsii, stem rot, white mould
jassids 434-7 359-60, 568, 591, 601, 606, 614,
lesser cornstalk borer 437, 445 645,693
soil insects 440 Screening for drought tolerance 310-12
southern corn rootworm 439 Screening methods for aflatoxin 530
Spodoptera litura 437-9 Scutellonema, yellow patch disease (S.
storage pests 462 cavenessi and S. clathricaudatum)
thrips 434-7 372,377
Resistance screening 558-9, 586 Seasonality of root growth 287-8
Responses (fertilizer) Secale cereale, rye 640
eu 238 Sections, genus Arachis
Fe 239, 240 Ambinervosae 98, 100, 108, 112, 119
K 230,231 Arachis 98, 100,102,105,107,109,113,
Mg231 119, 122
Mn240 Caulorhizae 98, 100, 103, 107, 108, 112,
P 228,229 119,633,645-55,657
Sn 233 Erectoides 98,100,104,107,108,112,
Zn 243,244 119
Reye's syndrome 527 Extranervosae 98, 100, 103, 109, 113, 119
RFLP 106, 575 Procumbensae 98, 100, 111, 119, 122
Rhizobia 226,227,260,624 Rhizomatosae 98, 100, 104, 107, 108, 113,
Rhizobium 260,624 126,633-45,654,657
Rhizoctonia solani 366-7, 369-70 Triseminalae 98, 100, 107, 112, 119
Rhizoma peanuts 633-5 Seed
Rhizomatosae (section) 98, 100, 102, 106, biochemistry 149-53
107, 108, 112, 118, 126, 633-45, biochemical evaluation 586
654, 657 borne nematodes 377-8
Rhodes grass (Chlorisgayana) 643 coat development 148-9
Rhopalosiphon padi 410 coat structure 86
Ricinus communis 374 deterioration 162
Riptortus serripes 426 development 144-9
Roasted peanuts 188 dormancy 153-155
Roasted peanut production 491-3 globulins 183
Root maturity indices 153
anatomy 76 metabolism 152-3
diseases 359-72 morphology 56-8, 125
growth 281 number per pod 73-4
hairs 60 proteins 105
hair invasion 256, 261 quality 158-62
knot nematodes 373-5, 649 storability and oil content 165-6
lesion nematodes 375-6 Seedling morphology 58
system 58-60 Selection for drought tolerance 310-17
Rust (Puccinia arachidis) 350-3 Semi-permanent cultivation 667-9
Rye (Secale cereale) 640 Sequential branching 63-4
Series in sections of genus Arachis
Salinity 274 section Arachis
Salted peanuts 492-4 series Amphiploides 98, 100, 107
Sampling methods (aflatoxin) 529, 530 series Annuae 98, 100, 107
Sanseveria libericum 460 series Perennes 98, 100, 107
732 Index
Series in sections of genus Arachis (contt£) Spodoptera ornithogalli 413
section Erectoides Spodoptera sunia 413
series Tetrafoliolatae 98, 100, 108, 119 Spotted wilt (tomato) TSWV 344-6
series Trifoliolatae 98, 100, 108, 119 Squalene 179
section Rhizomatosae Squirrels 577
series Prorhizomatosae 98,100,107, Sri Lanka 591,598
112, 119, 633 Stachyose 196-8
series Eurhizomatosae 98, 100, 107, 109, Standard crescent 68-9
112,633,636-7,639-41,643-5 Standard petal colour 68
Senegal 3, 4, 6, 576, 581-583 Starch 196-8
Serine 185, 186, 190 Stem
Setaria sphacelata var sericea 643 anatomy 76-8
Sethoxydim 642 colour 60-1
Shellers, hand operated 486-7 hairiness 61
Shellers, powered decorticators 487 morphology 60-4
Shelling 486 Stemphylium botryosum, melanosis 358, 645
Shifting cultivation 667-68 Stem rot, see Sclerotium rolfsii
Shoot growth 293 Stenomesius japonicus447
Single seed descent 574, 586 Sterol esters 178
Sitona 407 Sterols, free 178
Skimmed milk 195 Stocks, groundnut 15-17
Slow growing rhizobia 260 Stomata 78
Soil Storage 485-6
-borne pathogens 567-8 Storage pests 461-3
evaporation (Es) 302 Stripping 483-4
insects 399-400 Stylosanthes 99,107,258,624,653
nutrient depletion 632 Stylosanthinae (sub-tribe) 97, 99
preparation 460-1 Subspecies fastigiata 99, 123
water deficit, effects 305 Subspecies hypogaea 99, 123
water extraction 287 Subspecific classification 99, 123
Soluble carbohydrates 196-8 Sudan 5, 6, 576
Solvent extraction 188, 500 Sulphur (S) 214, 233-4
South Africa 5, 6, 584 Sunnhemp (Crotalaria juncea) 456
South-east Asia 3, 5, 6, 446-9, 601-6 Supplementary irrigation 317-8
Southern Africa,S, 6, 584--5 Survenola digitgrass, Digitariaxumfolozi 643
South-west Asia 3, 5, 6, 446-9, 591, Sympiesie dolichogaster 645
592-601 Synthetic mulches 457-8
Southern corn rootwork (Diabrotica Syrphids 449
undecimpunctata howardi) 439, 571 Systates articollis 406
Sowing times 680-3
Soybean (Glycine max) 559 Taiwan 3, 5, 6
Soy flour 195 Tamarindus indica 415
Spacings (plant) 680, 684--8 Tannins 203
Spanish (cultivar type) 55 Tanzania 5, 6, 576, 584
Specific, subspecific and subgeneric Taproot 58-9
nomenclature 97-8 Taxonomy 43 et seq., 97 et seq., 709, 710
Sphaceloma arachidis, peanut scab 653 Taylorilygns 412
Phenoptera indica, jewel beetle 402 Temik 567
Spiders 445 Temperature effect on seed quality 159
Spodoptera, army worms 413,571,574,577, Temperature, storage 485
586,591,598,601 Temperature tolerance 586
Spodoptera eridania 413 Tenebrionidae, false wireworms 406, 407,
Spodoptera exempta 413 468
Spodoptera exigua 413, 416, 468 Teratogenicity, aflatoxin 509
Spodoptera frugiperda 413, 416, 424--5, 468 Termites, Isoptera 399-400,414,468,571,
Spodoptera latifascia 413 577, 591, 692
Spodoptera littoralis 407, 413, 451 Tetrafoliatae (series) 98, 100, 109, 119,655
Spodoptera lilum 403,407,416,418,420, Tetrahymena pyriformis 191
424-6,446-7,464--5,468 Tetranychus 414
resistance 437-9 Tettigometridae 401
733
Textured vegetable protein (TVP) 193 Utilization
Thailand 3,5,6,446--9, 601,602,605 crop 718-19
Thanatephorus cucumenis, see Rhizoctonia forage 624 et seq.
solani germplasm 555
Therevidae 450 protein 191
Thiamine 200
Thielaviopsis basicola, blackhu1l363-4
Valencia cultivars 55
Thin layer chromatography (TLC) 531-2
Valine 186, 188, 193
Thiram 363, 462
Vapour pressure deficit 285
Threonine 183, 185, 186, 188, 190, 193, 194
Varietal classification 53-5
Thrips, Thysanoptera 344--6, 408, 427, 468,
Vegetative axis (V) 66
571,574,577,586,591,598,601,
Vegetavie growth 293
614,692
Venezuela 2, 4, 6
Thrips patmi 408, 409
Vernolate 642
Thrips tabaci 409
Verticillium dahliae 371-2
Thrips fusca 410
Verticillium wilt 371-2
Tocopherols 178-80, 200
Vertivera nigritana 460
Togo 3,4,6
Vicieae 267
Tolerance, pest 432
Vietnam 356, 446--9, 601, 605
Tomato spotted wilt virus (TSWV) 565,
Virginia cultivars 54
566, 586, 597
Viruses 339-46
Total biomass (W) 301
Vitamin A 179
Total dry matter (TDM) 304, 308
Vitamins 200
Total oil contact 175---{), 208
Vitamins, E, K, B groups 200
Toxicity
Vulgaris (var.) 55, 99, 553, 554
aflatoxin 509, 516, 520, 522, 523, 525-8
boron 234
manganese 241 Water
zinc 243-4 content 282
Toxin production (aflatoxin) on defined deficit effects 284-7
media 570 logging tolerance 586
Transpiration (T) 302, 306, 308 potential 282
Transpiration efficiency (TE) 308 relations 281 et seq.
Transport, calcium 215 resources 708-9
Trichogramma 445 stress 156--7, 159--61
Tridax procumbens 411 supply 665
Trifluralin 642 use efficiency 308
Trifolieae 267 West Africa 3, 4, 6. 576, 580--3
Trifoliolatae (series) 98, 100, 109, 119 Weed control 642
Trifolium 624 Wheat flour 194
Trifolium repens 642 White clover (Trifolium repens) 640
Triseminalae (section) 98,100,107,112, White fringed beetle (Graphognathus spp.)
119 571
Trypsin inhibitors 190, 196, 203 White grubs 402---{), 571, 577, 586, 591, 598,
Tryptophan 186--8, 190, 193, 200 692
Turgor potential 282 WHOIFAD reference protein 193
Turgor pressure 283 Wild Arachis species
Turkey 3, 4, 5 anatomy 79
Turkey X disease 509, 536 source of resistance 564, 570
Tylenchorhynchus 372, 375
Tylenchorhynchus brevilineatus 375, 376
Xanthomonas bacterial leaf spot 338-9
Tyrosine 185, 186
2, 4-DB 642 xylem sap analysis 269-70
Uganda 5, 7, 9,13 Yellow patch disease (Scutellonema

UNDP 576 cavenessi and S. clathricaudatum)
USA 2,4,6, 441---{), 608, 614 377
USAID 576 Yield, pods 306--8
USDA 614 Yield responses to evaporation/transpiration
Uruguay 2,4,6,613-14 305-9
734 Index
Yields, forage 649 Zimbabwe 5, 6, 576, 585
Ynchie 31 Zinc (Zn) 202, 233, 243
deficiency 243
Zaire 5, 6, 576, 584 toxicity 243
Zambia 576, 584, 585, 675-95 Zonate leaf spot (Cristulariella morieda) 358

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THE GROUNDNUT CROP

W orld Crop Series

IDIlI SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.

ISBN 978-94-010-4315-1 ISBN 978-94-011-0733-4 (eBook)

Library of Congrcss Catalog Card Number: 94-70983

§Printcd an acid-frec text papcr. manufacturcd in accordance with ANSII

1 Groundnut production and trade 1

2 The origin and history of the groundnut 24

3 Botany - morphology and anatomy 43

7 Mineral nutrition 214

11 Groundnut pests 395

12 Industrial utilization and processing 480

13 Mycotoxins in groundnuts, with special reference to aflatoxin 509

13.10 Overcoming aflatoxin contamination 535

K.J. Middleton (deceased)

Groundnut production and trade

This chapter focuses on major issues and implications of groundnut pro-

1.1 PRODUCTION, AREA HARVESTED AND YIELD

Several factors determine changes in yield, area harvested and level of

1.1.1 Groundnut production

(a) North America

(b) South America

1.1.2 Area harvested

Sudan 263.2 621.2 358.0 565.3 -55.9

reacted to improved production technology, changes in relative price,

(a) North America

Average 1960s: 1970s 1970s: 1980s

1-10 Pakistan (Brazil) USA (Pakistan)

area; however, an expansion of the groundnut export market and changes

(b) South America

Sudan 320 769 449 739 -30

Note Reported figures have been rounded to the nearest thousand.

third largest producer of groundnuts in Asia and its area harvested

Average 1960s:1970s 1970s:1980s

1-10 USA (Malawi) India (USA)

1.1.3 Groundnut yield

(a) North America

(b) South America

declined between 1960 and 1989 - in Ghana by 67% and in Cameroon by

Sudan 835 817 -18 753 -64

Average 1960s:1970s 1970s:1980s

1-10 Argentina (Japan) USA (Mexico)

1.2 STOCKS AND PRICES

Year US CN AG SD SAF GB ID BZ THL SEN VT MAL

1.3 INTERNATIONAL TRADE

1.3.1 International trade in groundnuts

1.3.2 International trade in groundnut oil

1.3.3 International trade in groundnut cake and meal

1.4 CONCLUDING COMMENTS

The author would like to thank D. Cummins, S. M. Fletcher and J. C.

The origin and history

Groundnut is a native New World crop. Early explorers found it culti-

Peruvian civilization apparently began along the eastern slopes of the

2.3 POST -COLUMBIAN HISTORICAL NARRATIVES

2.3.1 Sixteenth century natural histories

The first delibrate introductions of groundnut to Europe probably went

2.3.2 Selected seventeenth century narratives

2.S ORIGIN OF GROUNDNUT

2.S.1 Geographical origin of genus Arachis