Biology of Cajanus Cajan Pigeon Pea REVISED-1

Biology of Cajanus cajan (Pigeonpea)
Prepared by:
Ministry of Environment, Forest and Climate Change (MoEF&CC)
and Indian Institute of Pulses Research, Kanpur
under UNEP/GEF supported Phase II Capacity Building Project on Biosafety
Material from this publication may be used for educational purpose

provided due credit is given
Compiled by:
Dr. N. P. Singh Dr. S. K. Chaturvedi Dr. Dibendu Datta
Director, Indian Institute of Pulses Head (Division of Crop Improvement) Principal Scientist (Plant Biotechnology)
Research, Kanpur, Uttar Pradesh Indian Institute of Pulses Research, Indian Institute of Pulses Research,
Kanpur, Uttar Pradesh Kanpur, Uttar Pradesh
Dr. Alok Das Dr. Abhishek Bohra

Scientist (Plant Biotechnology) Scientist (Plant Breeding)
Indian Institute of Pulses Research, Indian Institute of Pulses Research,
Kanpur, Uttar Pradesh Kanpur, Uttar Pradesh
Reviewed by:
Dr. Ranjini Warrier Dr. O. P. Govila
National Project Coordinator, Former Professor of Genetics,
Phase II Capacity Building Project on Biosafety Indian Agriculture Research Institute
Advisor, MoEF&CC
Dr. Michael Wach Dr. Vibha Ahuja

Senior Scientist, Program Manager, Chief General Manager,
Centre for Environmental Biotech Consortium India Limited (BCIL)
Risk Assessment-ILSI Research Foundation
Consultation process:
The inputs and comments provided by the following institutions are gratefully acknowledged.
• National Bureau of Plant Genetic Resources, New Delhi
• International Crops Research Institute for the Semi-Arid Tropics, Hyderabad
• Banaras Hindu University, Banaras
• Vivekananda Parvatiya Krishi Anusandhan Sansthan, Almora
• Navasari Agricultural University, Navasari
• Rajmata Vijayrajsindhya Krishi Vishavvidhalya, Gwalior
• Dr. Balasaheb Swant Konkan Krishi Vidyapaeeth, Dapoli
• Tamil Nadu Agricultural University, Chennai
• University of Agriculture Sciences, Bangalore
Assisted in co-ordination & printing by:

Dr. P. Hareesh Chandra, Project Consultant and Ms Sonia Kaushik, Senior Project Executive, BCIL
Project Coordination Unit, Phase II Capacity Building Project on Biosafety
For further information, please contact:

Ministry of Environment, Forest and Climate Change
Government of India
Indira Paryavaran Bhawan, Jor Bagh Road, Aliganj
New Delhi 110003, Email: [email protected]
Contents
1. INTRODUCTION...................................................................................................... 1
1.1 Classification and Nomenclature ..................................................................... 1
1.2 Botanical Description of Pigeonpea ................................................................. 2
1.3 Economic Importance and Nutritional Composition....................................... 4
2. AREA, PRODUCTION AND PRODUCTIVITY........................................................ 5
2.1 Geographic Distribution.................................................................................. 5
2.2 Distribution in India including Regions of Cultivation ................................... 6
2.3 Zonalization and Varietal Testing System for Release........................................ 6
3. GEOGRAPHIC ORIGIN, GENOMIC EVOLUTION AND.................................... 7

CHROMOSOME NUMBER
3.1 Centres of Origin and Diversity....................................................................... 7
3.2 Genomic evolution.......................................................................................... 9
3.3 Genetic Diversity of Indian Germplasm........................................................... 10
3.4 Germplasm Collection at National and International Institutes........................ 11
4. REPRODUCTIVE BIOLOGY.................................................................................... 11
4.1 Reproduction................................................................................................... 11
4.2 Methods of Pollination, Known Pollinators and Pollen Viability...................... 11
4.3 Seed Production and Dispersal......................................................................... 12
4.4 Mating Systems................................................................................................ 12
4.5 Methods of Reproductive Isolation................................................................... 13
4.6 Potential of Vegetative Propagation.................................................................. 13

5. HYBRIDIZATION AND INTROGRESSION............................................................. 13
5.1 Intraspecific Crosses ........................................................................................ 13
5.2 Naturally Occurring Interspecific Crosses......................................................... 13
5.3 Experimental Interspecific Crosses.................................................................... 14
5.4 Genetic Introgression....................................................................................... 15
6. Human health implications.......................................................................... 16
7. Known interactions with other organisms in managed.............. 16

and unmanaged ecosystems
7.1 Interactions in Unmanaged and Managed Ecosystems...................................... 16
7.2 Important Insect Pests, Nature of Damage and their control in Managed......... 17
Ecosystems
7.3 Important Diseases, Causal agents and their Control in Managed Ecosystems.. 19
8. Agronomic Practices.................................................................................... 20
8.1 Climate............................................................................................................ 20
8.2 Sowing Season.................................................................................................. 20
8.3 Seed Production............................................................................................... 20
9. Breeding objectives......................................................................................... 22
9.1 Milestones in Breeding..................................................................................... 22
9.2 Heterosis Breeding in Pigeonpea ..................................................................... 22
9.3 Major Traits of Interest and Priorities in Breeding............................................ 22
9.4 Advancements and Challenges ......................................................................... 23
10. Biotechnological interventions in pigeonpea................................. 24
10.1 Transgenic Approaches for Genetic Improvement............................................. 24
10.2 Genomics and Molecular Breeding in Pigeonpea ............................................. 25
11. REFERENCES............................................................................................................ 27
Biology OF
Cajanus cajan L.
(PIGEONPEA)
1. INTRODUCTION The botanical name, C. cajan (L.) Millsp. has

been accepted universally for pigeonpea. The
1.1 Classification and taxonomic position (Van der Maesen 1990; http://
Nomenclature www.uniprot.org/taxonomy/3821) of pigeonpea
Pigeonpea [Cajanus cajan (L.) Millsp.] belongs [Cajanus cajan (L.) Millsp.] is as follows in Table 1:
to the genus-Cajanus, subtribe-Cajaninae, tribe- Table 1: Scientific Classification of Pigeonpea
Phaseoleae, order-Fabales,family-Fabaceae and Kingdom Plantae
sub-family Faboideae. Several edible beans like Division Magnoliophyta
Lablab, Dolichos, Phaseolus, Vigna and Cajanus Class Magnoliopsida
come under tribe Phaseoleae but in the sub-tribe Order Fabales
Cajaninae, only one species, Cajanus cajan has been
Family Fabaceae
domesticated and cultivated. The species belonging
Sub-family Faboideae
to Cajaninae have peculiar vesicular glands on
the leaves, calyx and pods which deposit asticky Tribe Phaseoleae
substance on their surface. It is the second most Subtribe Cajaninae

important pulse crop grown in India. Genus Cajanus
Species cajan
The term ‘pigeonpea’ was coined in Barbados,
where its seeds were considered an important De (1974) opined that differences between the two
pigeon-feed (Gowda et al., 2011). Pigeonpea or genera Atylosia and Cajanus are in size and vigour of
red gram or tur is known by several vernacular the plants, size and non-shattering of pods and size
names in India viz. Tur (Maharashtra and Gujarat), and number of seeds, which might have resulted
Arhar (Uttar Pradesh, Bihar, Madhya Pradesh), due to different forces during domestication.
Aral (West Bengal), Kandi (Andhra Pradesh), Roy and De (1965) and Van der Maesen (1981)
Harad (Haryana and some parts of western Uttar proposed the merger of the two genera (Atylosia and
Pradesh), Rahat (parts of Bihar), Tuvaraparippu Cajanus) because of their cytological similarity and
(Kerala), Kokh-lan (tribes of Tripura), adhaki and successful crossing with the diploid species. Van
tuvarika (Sanskrit). The alternate (Syn.) botanical der Maesen (1986) merged the species of Atylosia
names of pigeonpea are as follows: Cytisus cajan W. & A. with Cajanus DC on biosystematic
L.; C. bicolor DC.; C. flavus DC.; C. indicus grounds. Morphological, cytological, chemical
Spreng.; C. striatus Bojer (van der Maesen, 1990). and hybridization data support this merger, even
Biology of Cajanus cajan L. (PIGEONPEA) 1

if the required taxonomic changes are inconvenient sensitive short day plant and exhibit wide variation
(Van der Maesen, 1990). Van der Maesen (1990) in days to flower among genotypes (Gooding,
indicated C. ajanifolius as the most probable 1962; Spence and Williams, 1972).
progenitor of pigeonpea.
1.2.2 The Botanical Features of different plant
The genus Cajanus has 32 species, most of them
parts are as follows
being distributed in India and Australia. About
17 Cajanus species including the most probable Root
progenitor of pigeonpea [C. cajanifolius (Haines)
Pigeonpea has deep tap roots which extend
Maesen] occur in India. The variability available
vertically up to 2 meters and spread horizontally
within the cultivated species C. cajan forms the
through lateral roots. The Root is well developed
primary gene pool, while 11 species that are cross
in upper 60 cm soil profile (Natarajan and Willey,
compatible with C. cajan constitute the secondary
1980). The root proliferation is correlated with
gene pool. These cross-compatible species are
the duration of crop and growth habit (Sheldrake
C. acutifolius (F. Muell.) Maesen, C. albicans (Wight
and Narayanan, 1979; Mahta and Dave, 1931).
& Arn.) Maesen, C. cajanifolius (Haines) Maesen,
C. confertiflorus F. von Muell, C. lanceolatus (W.
Fitzg.) Maesen, C. latisepalus Maesen, C. lineatus
(Wight & Arn.) Maesen, C. sericeus (Baker) Maesen,
C. trinervius (DC) Maesen, C. scarabaeoides (L.)
Thouars Maesen, C. reticulatus (Dryand.) F. Muell.
These species are reported to be rich sources of various
desirable traits. The species viz. C. goensis Daiz,
C. heynei (W & A) Maesen, C. kerstingii Harms,
C. mollis (Benth) Maesen, C. platycarpus (Benth)
Maesen, C. rogosus (W & A) Maesen, C. volubilis
(Blanco) Blanco do not cross with the cultivated
pigeonpea and form the tertiary gene pool. Fig. 1: Piegonpea plant
1.2 Botanical Description of Stem

Pigeonpea An angular and woody stem originates from
three ribs starting from the base of each petiole.
1.2.1 Growth and development Starch present in xylem parenchyma and the
Pigeonpea is adapted to the tropical and subtropical medullary rays are mobilized to the pod and seed
(Sheldrake, 1984). Branching pattern (compact or
region and it can be grown on marginal land and
semi spreading or spreading) is determined by the
low fertilizer input, even under drought condition.
genetic constitution.
The growth habit is predominantly indeterminate
but some genotypes show determinate growth. The Pigeonpea plant show great plasticity by adjusting
branching pattern varies from erect to spreading. its branching behavior depending on the available
Pigeonpea is a predominantly photoperiod space between plants.

Leaves forms the standard; two lateral, obliquely obovate
and incurved clawed petals are known as wings; the
Leaves are spirally arranged, pinnately trifoliate and two innermost obtuse, incurved and boat shaped
lanceolate to oblong in shape. The terminal petiole petals are fused to form the keel to protect the
is highly variable and attains a length of 10-20 mm stigma and style. The standard and the wings are
while the lateral petiole is usually 2-3 mm long. generally of bright yellow colour, whereas keel is
Leaf size varies from 6-17 cm; lateral leaflets are greenish yellow. A lot of variation in petal colour
smaller than the terminal leaflet which varies from can be observed in the germplasm collections.
4-8 cm. Leaves are pubescent due to the presence The androecium has 10 stamens bunched into
of simple or glandular hairs (Bisen and Sheldrake, two groups (diadelphous) of 9 and a single free
1981). Figure 1 represents piegonpea plant. stamen that is attached at the base of androecium.
The grouped filaments are fused at the base and
1.2.3 Reproductive Parts cover the gynaecium, while the upper part is free
and bear uniform anther of about 1 mm length.
Inflorescence Six filaments are long, while the remaining four
The inflorescence is raceme which contain up to stamens including the free posterior have short
ten flowers per panicle and usually two flowers filaments which are supposed to encourage self
open at a time on a single inflorescence (Sharma fertilization (Bahadur and Rao, 1981). The
and Green, 1980). Flowering is acropetal (in the dorsifixed anthers, consisting of two halves, are
direction of apex), both within the raceme and pale yellow to yellow in colour. The placement of
on the branch. A single plant can hold up to 915 subsessile, dorsoventrally flattened, punctuates and
racemes (Remanandan et al., 1988). The terminal densely hairy ovary is superior. The long, filliform
or auxillary raceme is usually 4-12 cm long. In and glabrous style of gynaecium bears a thick,
most of the long duration genotypes the racemes incurved and capitate stigma. The short stalked
are grouped together at the end of branches, while glandular ovary is unilocular and monocarpellary
in early, medium and indeterminate genotypes the bearing 2-9 ovules with marginal placenta.
racemes are distributed along the branches (Sharma
and Green, 1980). Pod
Pod size is highly variable. The vegetable
Flower types have long pods with 4-7 seeds per pod.
The flowers are bisexual, zygomorphic and Depending on the genotype, 2-7 seeds develop
predominantly yellow (Sundaraj and Thulasidas, in each pod. Seeds are produced in separate
1980). More flowers are seen on the top of the locules and the pod may be highly constricted
peduncle. Small flowers, normally about 2 cm in certain genotypes giving beaded appearance.
in length are borne on thin, hairy pedicel. The Pod colour varies from green to dark purple with
flower size is very small in wild species. Flower varying degrees of brownish or purplish streaks.
size is correlated with seed size (Sharma and Pod is generally pubescent with varying degrees
Green, 1980). The calyx is gamosepalous with of simple or glandular hairs. Pod shattering at
five lobes. The corolla is zygomorphic and petals maturity is uncommon in cultivated varieties as
are imbricate. The largest, auricled and erect petal it is an undesirable trait for grain harvest.

Seed feed and fodder but also provide fuel wood for
the rural people, thus very popular among small
The germplasm of pigeonpea show a variety of seed
and marginal farmers. The dry sticks of pigeonpea
colour (white, creamy white, silvery, fawn, dark
plant are also used for making baskets, thatches and
purple which appear as black, pink, red to purple,
storage bins. In addition to atmospheric nitrogen
straw, brown) with or without specks and blotches
fixation through root nodulation by a wide range of
of different shades. The 100 gram seed weight
varies considerably from 5 to 22 g in germplasm symbiotic Rhizobia strains (Chikowo et al., 2004),
materials. The 100 seed weight of short duration the defoliated leaves also add nitrogen and organic
cultivated varieties are low (generally 6-8 grams) as matter to the soil (Mafongoya et al., 2006). Being
compared to long duration varieties (9-13 g). Seed a deep rooted legume, it also improves the physical
weight of medium duration varieties lie between condition of the soil for the next crop. Krauss
early and late maturing varieties. The 100 seed (1936) considered pigeonpea for soil binding
weight of vegetable types may reach up to 22 g. and advocated its plantation in Hawaii Island for
Seed do not show dormancy and germination is checking soil erosion.
hypogeal.
Crude protein ranges from 28–36% in green
1.3 Economic Importance and foliage of pigeonpea (Phatak et al., 1993) and
Nutritional Composition Table 2: Nutritional composition of mature pigeonpea seeds
Pigeonpea is one of the most important pulse crops Nutritional value per 100 g
of India. It is an integral part of the subsistence Energy 343 kcal
and rainfed farming systems in many parts of Carbohydrates 62.78 g
India. Being a hardy crop, it is a natural choice Dietary fiber 15 g
for small and marginal farmers particularly, in Fat 1.49 g
semi-arid dry-land areas because it can be grown Protein 21.7 g
successfully under rainfed or low input condition Thiamine (vit. B1) 0.643 mg (56%)
and provide nutritive food, feed, fodder and fuel Riboflavin (vit. B2) 0.187 mg (16%)
wood. In India, it is mainly consumed in the form Niacin (vit. B3) 2.965 mg (20%)
of split pulse as ‘dal’. Its immature green seeds Pantothenic acid (B5) 1.266 mg (25%)
and pods are also consumed as a green vegetable Vitamin B6 0.283 mg (22%)
by the tribal people of many States (provinces) Folate (vit. B9) 456 μg (114%)
such as Chattisgarh, Jharkhand, Andhra Pradesh, Calcium 130 mg (13%)
Karnataka, Gujarat and in the entire North-East Iron 5.23 mg (40%)
Hill region, where it is primarily grown in the Magnesium 183 mg (52%)
kitchen garden, backyards, hilly tract and on jhum Manganese 1.791 mg (85%)
land. The seed coat together with husk provides a Phosphorus 367 mg (52%)
valuable feed for milch animals. The green leaves Potassium 1392 mg (30%)
and tender branches provide nutritive fodder to Sodium 17 mg (1%)
livestocks. The tall and erect pigeonpea varieties Zinc 2.76 mg (29%)
are known to provide not only nutritious food, Source: Mazur et al., 1998

average seed protein content has been reported to is used in the traditional treatment of jaundice
be 21.7% (Mazur et al., 1998). Pigeonpea seeds and diabetes and used a antidote against food
provide essential amino acids like lysine, tyrosine, poisioning, gingivitis, stomatitis and constipation
and arginine, wheras cystine and methionine (Lans, 2007; Ganeshan, 2008; Upadhyay et
contents are low (Saxena et al., 2010a). Pigeonpea al., 2010). Several anti-nutritional factors like
seeds are also rich in potassium, phosphorus, trypsin inhibitor, protease inhibitors, amylase
magnesium, calcium and iodine. However, cystine inhibitors, cyanogenic glycoside, hemagglutinin,
and methionine contents are low (Nwokolo, alkaloids and tanninphytolectins, polyphenols and
1987). It also contains fair amount of iron and oligosaccharides have been reported in pigeonpea
selenium and small amount of zinc, copper and seeds. However, simple processing methods like
manganese, vitamin A, niacin and small amount soaking in water, boiling, cooking, germination
of thiamine, riboflavin, vitamin B6, folate and and fermentation reduce the level of these factors
pantothenic acid (Table 2). Pigeonpea leaf extract (Singh, 1998; Onwuka, 2006).
2. Area, production and productivity

2.1 Geographic Distribution been introduced in China as well where it is planted
Distribution of pigeonpea is asymmetric over the on the hilly slopes primarily to check soil erosion
globe. It is grown in different parts of the world (Saxena, 2008).
covering more than 22 countries including India, Globally 4.33 million tonnes (mt) of pigeonpea
Myanmar, Tanzania, Malawi and Kenya (Fig 2) was produced and India alone contributed 2.65
(FAOSTAT, 2013). In South-East Asia, pigeonpea mt followed by Mayanmar (0.9 mt), Tanzania
is mainly grown in India, Mayanmar, Nepal, (0.3 mt), Malawi (0.24 mt), Kenya (0.09 mt) and
Bangladesh and Phillipines. It is widely cultivated Uganda (0.084 mt) (FAO STAT, 2012). The area
in India where it plays an important role in pulse and production of pigeonpea for last 6 years in
based cropping systems and occupies second largest India has been presented in Table 3.
area among the pulse crops. Recently this crop has
Table 3: Area and Production of Pigeonpea in India
Area Production
Year
(Million hectares) (million tonnes)
2007-08 3.73 3.08
2008-09 3.38 2.27
2009-10 3.53 2.46
2010-11 4.42 2.86
2011-12 4.04 2.65
2012-13 3.81 3.07
2013-14 3.88 3.29
Source: Agricultural Statistics Division, Directorate of
Fig. 2: Major global producers of pigeonpea (%) Ecomomics and Statistics, Department of Agriculture
Source: FAOSTAT, 2013 and Co-operation, Ministry of Agriculture.

2.2 Distribution in India including cultivated species of pigeonpea does not exist as
Regions of Cultivation naturalized population in the wild form in any
ecological zone of India. Hence its natural habitat
India occupies the largest area (3.5 – 4.0 million conditions are not known; but it prefers grassy
hectares) of pigeonpea in the world, therby habitats in tropical, cold free zones with optimum
contributing nearly 80% area globally. Although 600-1000 mm annual rainfall. Natural population
pigeonpea is grown in 315 districts of India, 26 of various wild species of pigeonpea can be found in
districts account for about 50% area (Bhatia et al., Eastern and Western Ghats, North-Eastern states
2006). In India, it is a widely grown crop covering and in forests and hilly areas in almost every state
more than 18 states. About 85% of the pigeonpea (Sardana et al., 2011). Generally, wild species of
is grown in six states namely, Maharashtra, pigeonpea are spotted on the sunny and drained
Karnataka, Madhya Pradesh, Andhra Pradesh, area of the forest edges, in open places within the
Gujarat and Jharkhand (Fig.3). Other states forest or on grasslands and hill slopes.
include Uttar Pradesh, Orissa, Tamil Nadu, Bihar
and Chattisgarh. To a limited extent, pigeonpea 2.3 Zonalization and Varietal
is also cultivated in Rajasthan, Punjab, Haryana,
Testing System for Release
West Bengal and North-Eastern States. Pigeonpea
is grown traditionally in the foot hill regions of
The Project Directorate on Pulses (PDP), Kanpur
Dehradun upto 1500 m altitude, where it is locally
Uttar Pradesh was created to look after the All
known as ‘Tur’.
India Co-ordinated Project on Improvement of
Pulses (AICPIP) including pigeonpea. The PDP
was later upgraded to the level of Directorate of
Pulses Research (DPR) in 1984 and subsequently
in 1993, it attained the status of Indian Institute
of Pulses Research (IIPR). The AICPIP was tri-
furcated, and a separate network, All India Co-
ordinated Research Project (AICRP) on Pigeonpea
was created for zone based co-ordinated research
catering to the specific needs of pigeonpea.
Pigeonpea is a photo thermo sensitive crop and

Fig. 3: Major state wise producer of pigeopea (%) its phenology differs with the climatic conditions
[Source: GoI, Department of Agriculture & and varieties. Long duration varieties (>180 days)
Cooperation (2014-15)]
are pre-dominant in North East plains (Eastern
In India, pigeonpea crop has four distinct maturity Uttar Pradesh, Bihar, Jharkhand, West Bengal,
groups viz., early (120-140 days), mid-early (141- Assam) and parts of Northern Chattisgarh and
160 days), medium duration 161-180 days) and Bundelkhand region of Uttar Pradesh and Madhya
long duration (>180 days). Although pigeonpea Pradesh. Medium duration (161-180 days) varieties
can grow well on hilly slopes, grass lands, forest are usually grown in Central (Chhatisgarh, Madhya
lands, degraded lands and ravine areas the Pradesh, Gujarat, Maharshtra) and Southern India

(Andhra Pradesh, Karnataka, Tamil Nadu and parts
of Odisha), whereas early duration varieties are pre-
dominant in North West plains (Punjab, Haryana,
parts of Rajasthan and western Uttar Pradesh).
For the purpose of varietal release, elite pigeonpea

breeding lines are evaluated to assess their
performance with respect to grain yield, disease
resistance and their suitability for cultivation
in specific zone(s). Testing centres and growing
areas have been dividied into five zones (Fig 4),
namely-
1. North Hill Zone (NHZ) comprising

Uttarakhand, Tripura, Nagaland and Assam
2. North East Plain Zone (NEPZ) comprising
central and eastern Uttar Pradesh, Bihar, Fig. 4: Zonalization and testing centres of pigeonpea
Jharkhand and parts of West Bengal
3. North West Plain Zone (NWPZ) comprising They are also involved in technology transfer and
Delhi, Haryana, Rajasthan,Punjab, western quality seed production and are catering to specific
Uttar Pradesh and tarai region of Uttarakhand needs of various agro-climatic zones. There is a three
tier varieties testing system for release. Promising
4. Central Zone (CZ) comprising Madhya
elite breeding lines are evaluated as entry in Initiatial
Pradesh, central and southern Chattisgarh,
Varietal Trial (IVT) across the zone(s), and the
Gujrat and Maharashtra
genotype(s) performing better than check variety
5. South Zone (SZ) comprising Orissa, are promoted for further evaluation in Advanced
Telangana, Andhra Pradesh, Karnataka and Varietal Trial 1 (AVT 1) and Advanced Varietal
Tamil Nadu Trial 2 (AVT 2) on the basis of performance in
These zones have manpower and resources to a specific zone or zones. The elite breeding lines
evaluate elite breeding lines and participate in are recommended for release on the basis of
development of matching crop production and performance (3 years or more) in multi-locational
protection technologies. trials in comparision to the best check variety.
3. Geographic origin, genomic evolution and

chromosome number
3.1 Centres of Origin and Diversity including occurrence of various wild relatives
(Table 4) in nature, vast genetic variability in the
India is considered as the centre of origin for gene pool, and a few historical as well as archa-
pigeonpea (Vavilov, 1951). Many evidences eological records have been offered to strengthen

the view of Indian origin of pigeonpea (Van der Kassa et al., 2012). Fifteen wild species have been
Maesen, 1986, 1990). The alternate hypothesis reported in Australia also. Noticeably, majority of
suggesting Africa as the centre of origin does these species are endemic, therefore, Australia is
not seem to be viable as only one wild relative considered as the centre of diversity for pigeonpea.
C. kerstingii is reported to occur in West Africa. However, Kassa et al. (2012) contest this view that
In addition, C. scarabaeoides has also been found considers Australia as the centre of diversity since
in Africa, but spread is restricted to the coastal they observed very low level of genetic diversity
areas only. Consequently, van der Maesen (1980) among the wild Australian species, which were used
proposed Africa as the secondary centre of origin for SNP based genetic diversity analysis.
(Saxena, 2005). Therefore, the most acceptable
route of dispersion describes that the immigrants C. cajan having a pantropical distribution is the only
moved the crop up from India to East Africa, then cultivated species belonging to the genus Cajanus.
the route followed to Egypt (via Nile valley), West Pigeonpea underwent domestication around
Africa and finally to the America (Odeny, 2007a; 3,500 years ago (Vavilov, 1951; De, 1974; Royes,
Table 4: Wild Species and their Distribution in India
Species Distribution Reference(s)/Links
C. scarabaeoides Widely distributed species across Upadhyaya et al., 2013
Andaman and Nicobar, Andhra Pradesh,
Bihar, Chattisgarh, Himachal Pradesh,
Jharkhand, Karnataka, Kerala, Madhya
Pradesh, Maharashtra, Orissa, Tamil
Nadu, Uttar Pradesh, Uttaranchal,
West Bengal
C. albicans Peninsular India van der Maesen, 1990
C. cajanifolius Andhra Pradesh, Arunachal Pradesh, http://www.iucnredlist.org/details/full/19891613/0
Assam, Bihar, Chattisgarh, Gujarat,
Haryana, Himachal Pradesh, Jammu-
Kashmir, Karnataka, Kerala, Maha-
rashtra, Manipur, Meghalaya, Mizoram,
Nagaland, Orissa, Pondicherry, Punjab,
Rajasthan, Sikkim, Tamil Nadu,
Tripura, Uttar Pradesh, West Bengal
C. crassus (King) Maesen Assam, Central Indiaand NW Himalaya van der Maesen, 1990
var crassus
C. elonagatus (Bentham) NE India van der Maesen, 1990
Maesen
C. grandiflorus (Bentham ex Himalayn region van der Maesen, 1990
Baker) Maesen
C. mollis Himalayn region van der Maesen, 1990
C. platycarpus Bihar, Gujarat , Haryana, Himachal http://www.legumes-online.net/ildis/aweb/td114/
Pradesh, Jammu-Kashmir, Madhaya td_24020.htm
Pradesh; Maharashtra, Orissa, Punjab,
Rajasthan and Uttar Pradesh
(Source: http://www.theplantlist.org)

1976). Based on the morphological evidences, C. and Obermayer, 1998; Ohri and Singh, 2002).
cajanifolius is considered as the putative progenitor
of present cultivated pigeonpea (De, 1974; van The comprehensive karyotype analysis reported
der Maesen, 1986,1990). Morphologically, by Ohri and Singh (2002) included cultivars of
C. cajan and C. cajanifolius share similar attributes C. cajan and other wild species belonging to
except for the strophiole characteristics (De, 1974). Cajanus, Rhynchosia, Dunbaria, Flemingia
An elaborated comparison between C. cajan and and Paracalyx and they also reported the 4C
C. cajanifolius was established based on various DNA content in these species. With regard to
morphological attributes, extent of crossability chromosome structures, variations were detected
and cytology of the derived inter-specific hybrids in chromosome length and satellite choromosomes
(Mallikarjuna et al., 2012). (Naithani, 1941; Sinha and Kumar, 1979; Sharma
and Gupta, 1982; Pundir and Singh, 1986).
3.2 Genomic Evolution The drafts of pigeonpea whole genome sequence
Pundir and Singh (1985) reviewed the evolution yielded valuable insights about the genome
of C. cajan and proposed that an inter-specific evolution of pigeonpea (Varshney et al., 2011;
hybridization event between C. scarabeaoides and Singh et al., 2011). Based on K-mer statistics the
C. cajanifolius led to the evolution of C. cajan. entire genome size of pigeonpea was estimated to
Further, using PCR-RFLP technique different be 833.07 Mb. Furthermore, the whole genome
genera belonging to the sub tribe Cajaninae viz., assembly of pigeonpea also supports existence of 11
Cajanus, Rhynchosia, Dunbaria, Flemingia and pairs of homologous chromosomes. The genome
Paracalyx were studied and involvement of C. shed new light into the evolutionary dynamics
cajanifolius as the maternal parent was advocated of pigeonpea and absence of recent genome wide
(Lakshmi et al., 2000). duplication events was speculated by the authors
(Varshney et al., 2011).
The first report demonstrating the eleven pairs of
homologous chromosomes (n=11) in pigeonpea Based on the variation in esterase enzyme examined
was documented by Roy (1933). Afterward, an in the seed extracts of C. cajan (T21) and six wild
investigation using pollen mother cells strengthened species, Krishna and Reddy (1982) postulated
the hypothesis of haploid choromosome number that C. cajanifolius is the closest wild relative of
to be n=11 (Krishnaswamy and Ayyangar, 1935). C. cajan. Later, Kollipara et al. (1994) investigated
Somatic chromosome number of pigeonpea was the electrophoretic migration patterns in trypsin
reported to be 2n=22 (Naithani, 1941). The and chymo-trypsin inhibitors among 69 strains
wild relatives of pigeonpea also contain similar of C. cajan and 17 accessions belonging to seven
number of chromosomes except in African different wild Cajanus species. Similarly, close
species C. kertsingii, which exhibited a different proximity of C. cajan with C. cajanifolius was also
chromosome count of 2n = 32 (Gill and Hussaini demonstrated by Jha and Ohri (1996) through
1986). Thenceforth, a series of karyotype studies developing seed protein profiles of different
and measurement of the 4C DNA content were cultivated and wild accessions. Furthermore, the
done in pigeonpea (Ohri et al., 1994; Greilhuber karyotype analysis and variation in nuclear DNA

content revealed that the karyotype including number of primary branches, branching
morphology and number of satellite chromosomes length, crop duration, photoperiod sensitivity,
of C. cajan was the most similar to that of number of pods per plant, pod length, number
C. cajanifolius (Ohri and Singh, 2002). Several of seeds per pod, flower number, flower
other studies, particularly the molecular marker- colour, inflorescence size, pod colour and seed
based diversity analyses, supported C. cajanifolius size, seed color, taste, pod dehiscence, seed
as the most probable progenitor sharing a dormancy, seedling vigour, habitat preferences
similar pattern of DNA variation with C. cajan. and biochemical constitution.
Nadimplalli et al., (1993) used RFLP markers
to examine the relationship between 24 diverse Most of the pigeonpea genotypes are photo-
accessions belonging to four different genera, and thermo sensitive and a few behave like photoperiod
they esbalished a close relationship between C. insensitive. Therefore, days to 50% flowering and
cajan and C. cajanifolius. Higher genetic similarities physiological maturity do not correspond across
between C. cajan and C. cajanifolius were also the North-South region. Grossly, as mentioned
revealed by other marker systems including earlier Indian pigeonpea genotypes have been
mitochondrial DNA-RFLP (Sivaramakrishnan et classified into distinct maturity groups i.e. early,
al., 2002) and genomic SSR markers (Odeny et al., mid-early, medium and late. Vegetable types (tall,
2007b; Saxena et al., 2010c). More recently, Kassa large flower, bigger pods and seed) are distributed
et al. (2012) performed a comprehensive diversity in the North East Hill region, Gujarat, Karnataka,
analysis using over 700 SNP markers across 110 Chhattisgarh and Andhra Pradesh. Genotypes with
accessions (79 cultivated and 31 wild), and the non-branching habit (single stem) have also been
experimental evidences supported the hypothesis identified among collections from Madhya Pradesh
of C. cajanifolius being the closest wild progenitor and other parts of Central and Northern India.
of cultivated pigeonpea. Similarly by analyzing Large amount of variability exists for growth habit
1,616 SNP markers in a panel of 184 pigeonpea (determinate and non-determinate) and branching
accessions (77 cultivated and 107 wild), a close pattern (erect to spreading). Flower colour is
relationship of C. cajan was observed with C. predominantly yellow among the accessions of grain
cajanifoilus as compared to the other wild relatives type, while purple flower is predominant among
(Saxena et al., 2014). the vegetable types collected from Chhattisgarh
and Tripura. Wide scale variation is observed for
3.3 Genetic Diversity of Indian seed size and seed colour among the germplasm
collections. Considerable diversity was observed
Germplasm
in pigeonpea germplasm accessions of Bastar
Pigeonpea is grown in differtent agroclimatic region (Nag and Sharma, 2012). Forty nine
regions of India on a variety of soil and diverse pigeonpea genotypes originating from different eco-
physiographic situations. Wide range of varia- geographical regions exhibited considerable genetic
bility has been documented for various botanical diversity for 12 characters (Rekha et al., 2011).
and agronomic traits viz. plant growth habit, Variability for SSR alleles was shown among 36
plant height, days to 50% flowering, days to pigeonpea lines with varying degrees of resistance
physiological maturity, branching pattern, against Fusarium wilt (Singh et al., 2013).

3.4 Germplasm Collection at has been grouped from the core collection and other
National and International materials. In the national gene bank at National
Institutes Bureau of Plant Genetic Resources (NBPGR),
10,189 accessions of Indian collections are conserved.
There are 13,771 accessions of pigeonpea conserved In addition, NBPGR has also repatriated 5,748
at ICRISAT (Upadahyay et al., 2007). ICRISAT accessions to national gene bank. The extensive use of
has developed a composite collection of pigeonpea few parents in pigeonpea improvement programmes
containing 1000 accessions representative of the has led to the narrowing down of genetic base of
diversity of all germplasm collection. Furthermore, the cultivated varieties which defeats the purpose of
a mini core collection comprising 146 accessions collecting a large number of germplasm.
4. Reproductive Biology
4.1 Reproduction the middle of the flowering branch and then move,
in either directions. The flowers are papileonaceous
The basic floral arrangement in pigeonpea is
(completely bisexual and zygomorphic). Generally
typical of fabaceae family exhibiting terminal or
the stigma of a mature flower bud is surrounded
auxiliary raceme inflorescence. The panicles are
by anthers which dehisce a day before the opening
either terminal, as in the case of indeterminate
of flower (Fig. 5). On a bright sunny day, anthesis
types, or corymb-shape bunch in the determinate
starts in the early morning, peaks at 9-10 AM
types. The inflorescence has a long peduncle and
and continues till 4 PM. The duration of flower
flowers are concentrated at the end of branches in
opening varies from 6 to 36 hours depending upon
late maturing and determinate genotypes, whereas
the climate and environmental conditions.
flowers are borne along the branches in most of
the early, medium and non-determinate genotypes
(Sharma and Green, 1980). In general, flowering
4.2 Methods of Pollination, Known
starts from the base and proceeds acropetally Pollinators and Pollen Viability
towards the apex both within the raceme and on Pigeonpea possesses cleistogamous flowers which
the branch. In some cases the first flower appears in favour self pollination. However, 14-20 % natural
outcrossing was observed in pigeonpea (Sharma
and Green, 1980; Howard et al., 1919). Pigeonpea
is often-cross pollinated through entomophily. Self-
pollination occurs in the bud before the flowers
open, while cross pollination is effected with the
help of insects. Reddy and Mishra (1981) reported
low frequency of self fertilization when flower
buds were pollinated with foreign pollen without
emasculation. Onim (1981a) observed that
anthers dehisce during the bud stage but they start
Fig. 5: Open flowers in pigeonpea germinating 24-28 hours after dehiscence when

flowers start to wither. Pigeonpea is protogynous shattering. Pigeonpea seeds do not show dormancy
and the stigma becomes receptive 68 hours before and can be grown immediately after harvest
anthesis and stigma receptivity is maintained (Andersson and de Vicente, 2010).
even 20 hours post anthesis (Prasad et al., 1977).
Fertilization occurs within 48-54 hours after 4.4 Mating Systems
pollination (Dutta and Deb, 1970). Several factors
Large and bright coloured flowers coupled with the
like flowering habit of a genotype, presence of insect
presence of nectar attract a variety of pollinators.
population, temperature, humidity, wind velocity
On an average 20% out crossing has been observed
and wind direction affect natural outcrossing at a
due to insects in pigeonpea (Saxena et al., 1990).
given time and space (Bhatia et al., 1981). Most
To harness the potential of outcrossing nature
important pollinators are Apis spp. (A.dorsata, for heterosis breeding, instances of genetic male
A.laboriosa, A. florae and A. cerana), Megachile spp. sterility (GMS) systems were reported in pigeonpea
(M. lanata and M. flavipes) and Xylocopa spp. (Pathak, which was achieved by spontaneous mutations
1970; Williams, 1977). The rate of outcrossing (Saxena et al., 2010b). GMS plants are eliminated
varies from place to place depending on the extent of from the population unless they outcross with
pollinator bees and climatic conditions. Outcrossing male fertile genotypes. Male sterile plants induced
ranged from 20 to 70 % at various locations (Saxena through mutagen could not be maintained further
et al., 1990, 2010b). High rate of out crossing in (Dundas, 1990). Recently, cytoplasmic-genetic
pigeonpea creates problems in the maintenance of male sterility (CGMS) system has been established
varietal purity. in pigeonpea based on a range of sterilizing
cytolplasms viz., A1 cytoplasm derived from
4.3 Seed Production and Dispersal (C. sericeus), A2 cytoplasm from C. scarabaeoides,
Majority of flowers drop before developing into a A3 cytoplasm from C. volubilis, A4 cytoplasm from
pod and only a small portion gives rise to mature C. cajanifolius, A5 cytoplasm from C. cajan, A6
pods (Datta and Deb, 1970; Howard et al., 1919). cytoplasm from C. lineatus, A7 cytoplasm from C.
There is rapid development of endosperm during platycarpus (Saxena et al., 2010b). Figure No. 6a
the first week after fertilization. Seed development shows the visible difference between the anthers of
is visible 7 days after pollination. Though the seed sterile and fertile flowers.
of 30 days old pod attains physiological maturity,
it requires 10-15 additional days for obtaining seed
with desirable moisture content. Pod shattering is
the natural means for seed dissemination in wild
relatives of pigeonpea and these seeds germinate
in next crop season. Therefore, restriction of two
years has to be made to grow pigeonpea in the
same fields/plots. The pods of cultivated pigeonpea
are by and large indehiscent and do not shatter at
maturity. However, if the mature plants are left in Fig. 6a: Difference in anthers and pollen load in fertile
the field for a long time, pods show tendency of and sterile flowers

4.5 Methods of Reproductive
Isolation
Reproductive isolation is essential for the
production of genetically pure seeds as pigeonpea
is often cross pollinated crop and therefore.
Depending on the purity standards, a minimum
of 200 m (for certified seed) to 500 m (for breeder
seed production) isolation distance is required
Fig. 6 b: A scheme illustrating production of CGMS-based for producing true to type seed of a genotype.
hybrids and maintenance of parental lines Alternatively, entire seed production plot may
be isolated using appropriate nets which is rarely
CGMS approach involves three lines i) A-line
feasible under normal circumstances unless it is
ii) B-line which is isogenic to A-line except for
necessary for certain specific programmes like
the cytoplasm and iii) R-line that should restore
maintenance of CMS lines in hybrid breeding. The
fertility in the F1 hybrids. Based on their specific
repoductive isolation can be maintained by means
utility in the CGMS programme, A, B and R
of bagging the individual plant with nylon nets
lines are also referred to as sterile-line, maintainer-
before flowers initaition i.e. at flower bud initiation
line and restorer-line, respectively. Owing to
(Saxena, 2006; Saxena and Nadarajan, 2010).
the involvement of three different lines, CGMS
approach is also known as ABR system. Technically,
4.6 Potential of Vegetative
the CGMS breeding scheme comprises of two
Propagation
components that is production of F1 hybrids
and maintenance of parental lines. As illustrated Pigeonpea plants can be regenerated from the
in Figure 6b, the fertile hybrids are produced by vegetative tissue in the artificial medium (Chandra
crossing A-line with R-line i.e. ‘A × R’ progeny. and Pental, 2003; Thu et al., 2003; Eapen, 2008).
Concerning maintenance of A-line, it is crossed Like other grain legumes, seed is the key source of
with its isogenic line i.e. B-line. On the other hand, propagation. However, regrowth has been noticed
R and B-lines are maintained simply by selfing the in pigeonpea after cutting or ratooning the plants
individual plant. (Andersson and de Vicente, 2010).
5. Hybridization and introgression
5.1 Intraspecific Crosses Kumar, 2010), A. dorsata (Williams, 1977) and M.

velutina and Xylocopa sp. (Li et al., 2011).
Pigeonpea is an often cross pollinated crop in nature,
the extent of natural open pollination goes upto
5.2 Naturally Occurring
24% (Bhatia et al., 1981). The natural pollination
Interspecific Crosses
is usually mediated through a variety of insects
particularly Megachile lanata, Apis florea (Pathak, Based on the extent of crossability or gene flow,
1970), M. conjuneta and M. bicolor (Saxena and the concept of gene pool was proposed by Harlan

and de Wet (1971). A total of 31 species of (10%), C. sericeus (2.3%) and C. lineatus (17%)
pigeonpea are distributed across primary (only compared to the pigeonpea line (ICP 5529),
one, C. cajan), secondary (10 species) and tertiary which exhibited 22% out crossing. Notably, the
(20 species) gene pools (Ramanadam, 1990). low levels of out crossing observed in the wild
Figure 7 depicts distribution of these species across species may be attributable to the non-preference
different gene pools. of pollinating insects discarding the possibilities of
existence of potent compatibility barriers. Several
species belonging to secondary gene pool exhibit
sexually compatible reactions with C. cajan include
C. cajanifolius (Reddy et al., 1981; Pundir
and Singh, 1987; Yadav and Padmaja, 2002),
C. scarabaeoides (Roy and De, 1965; Pundir and
Singh, 1987), C. albicans (Pundir and Singh,
1987), C. sericeus (Kumar et al., 1985; Pundir
and Singh, 1987; Yadav and Padmaja, 2002),
C. lanceolatus and C. latisepalus (Kumar, 1985),
C. reticulatus and C. acutifolius (Dundas, 1984;
Reddy et al., 2001; Mallikarjuna and Saxena,
2005).
Fig. 7: A diagram illustrating the pigeonpea gene pool 5.3 Experimental Interspecific
(Sources: Mallikarjuna et al., 2011; http://ksiconnect. Crosses
icrisat.org/wp-content/uploads/2013/03/conquering-
gene-pools-pigeonpea.revised.pdf ) In general, the species belonging to secondary
gene pool are easily crossable with the cultivated
pigeonpea using the traditional hybridization
In regard to the natural inter-specific hybridization,
methods. This in turn avoids the need for
several instances of out crossing leading to the employing additional technical interventions
development of viable inter-specific hybrids including hormone-aided pollination and embryo
have been documented in pigeonpea (Saxena rescue. However, these interventions may be
and Kumar, 2010). Saxena and Kumar (2010) required to enable recovery of greater quantities
examined the degree of natural out crossing in of the hybrid seeds from the interspecific cross
the four wild relatives from the secondary gene (Mallikarjuna et al., 2011). However, reciprocal
pool viz. C. scarabaeoides (ICPW 89), C. albicans crosses involving wild species as female parent
(ICPW 13), C. sericeus (ICPW 162) and C. lineatus did not produce healthy embryos (Mallikarjuna
(ICPW 42). The percentage of natural out-crossing and Saxena, 2002). Thiruvengadam and Muthiah
was calculated on the basis of the number of hybrid (2007) generated viable hybrids from the cross
individuals observed in the succeeding generations. C. cajan × C. cajanifolius only when they used
Consequently, variable levels of out-crossing were C. cajanifolius as male parents, while reciprocal
reported like C. scarabaeoides (8.3%), C. albicans crosses yielded hybrids that could not set

pods. Similarly, crosses between C. cajan and 1995). However, while investigating inter-specific
C. acutifolius provided aborted embryos when hybridization using another species from tertiary
C. acutifolius was chosen as female parent gene pool (C. volubilis × C. cajan) existence of pre-
(Mallikarjuna and Saxena, 2002). Concerning fertilization barriers was also reported (Jayaprakash
cytology, while characterizing the hybrids (Cajanus and Sarla, 2001) and application of ε-amino
cajan × C. reticulatus var. grandifolius), it was caproic acid (EACA - an amino acid) to the pollen
observed that during diakinesis and metaphase-I, germination medium was suggested as it resulted
only 41.9 % of the cell had bivalents. On the in better pollen germination. To address the issue
other hand, several abnormal forms including of early embryo abortion, application of gibberellic
quadrivalents, trivalents, and univalents were acid (GA3) to the pollinated pistils has been
detected to considerable extent i.e. upto 45.4% advocated to allow recovery of mature embryos
(Reddy et al., 2001). that are usually more responsive to embryo rescue
technique (Mallikarjuna et al., 2011). In addition,
In contrast to compatibility with secondary gene application of GA3 or kinetin is also known to
pool, regular abortion of hybrid embryos has been enhance the rate of pod set and number of seeds
reported in case of experimental crosses involving per pod during inter-specific hybridization (Kumar
parents from tertiary gene pool in particular C. et al., 1985; Dhanju and Gill, 1985).
platycarpus (Pundir and Singh, 1987). However,
to assist inter-specific hybridization with C. 5.4 Genetic Introgression
platycarpus, effective techniques involving
Wild relatives of pigeonpea are known to contain
colchicine treatment and embryo rescue using
several agriculturally important genes that impart
in vitro culture practices have successfully been
tolerance to a wide range of biotic and abiotic
employed in pigeonpea (Mallikarjuna and Moss,
stresses such as sterility mosaic disease (SMD),
1995; Mallikarjuna, 1998; Mallikarjuna et
phytophthora blight, root-knot nematode, pod
al., 2006). By using embryo rescue technique,
borer, pod fly, salinity and drought etc. (Rao et
Mallikarjuna (2007) successfully generated several al., 2003; Bohra et al., 2010). In this context,
BC4F1 plants from the cross (C. platycarpus × distant hybridization emerges as a viable option
C. cajan) × C. cajan. Alternatively, to escape the not only to incorporate beneficial exotic alleles
cumbersome embryo rescue practises, colchicine of desired traits into cultivated-types, but also
treatment of the derived F1 (C. platycarpus × for broadening the extremely narrow genetic
C. cajan) plants was suggested by Mallikarjuna base of pigeonpea. In terms of trait-introgression,
et al. (2011), who developed tetraploid plants inter-specific hybridization led to the recovery
showing enormous vegetative growth. However, of remarkably distinct phenotypes in pigeonpea.
variations in ploidy levels did not permit further For example, some of the derivatives of the wild
back crossing of these tetraploid individuals (4×) cross (C. cajan × C. scarabaeoides) exhibited
with the cultivated C. cajan (2×). The occurrence enhanced level of protein content i.e. more than
of post-zygotic barriers has been proposed as the 27% (Reddy et al., 1997). Besides higher protein
underlying reason for the incompatible response content, Reddy (1990) obtained a valuable
mentioned above (Mallikarjuna and Moss, segregant with different plant architecture (dwarf

stature, designated as D0) from an inter-specific cross-pollination were also recovered from another
cross involving C. cajan and C. scarabaeoides inter-specific cross i.e. C. cajan × C. lineatus
as parents. Similarly, progenies with partially (Saxena et al., 1998).
cleistogamous flowers showing very low level of
6. human health implications

Pigeonpea seeds are known to contain 18-26 % case of kidney related disorders. Similarly, leaf
protein and in case of wild Cajanus species up to extracts from pigeonpea are reported to have
30% protein content has been observed (Odeny, noticeable impact in curing diverse diseases such
2007a). The nutritional value of pigeonpea is as malaria, diabetes, dysentery and hepatitis
evident from the information provided in Table (Oke, 2014). The flavonoids found in pigeonpea
1. Additionally, Saxena et al. (2010a) have leaves have important pharmacological properties
enlisted a range of major anti-nutritional factors such as anti-cancer and anti-inflammatory
which are reported in pigeonpea. These anti- (Saxena et al., 2010a). Similarly, the anti-cancer
nutritional factors include phenolic compounds, properties of cajanol (an isoflavanone isolated
tannins, phytolectins, oligosaccarides like raffinose from pigeonpea roots) were also demonstrated in
and stachyose and a variety of inhibitors that vitro against human breast cancer cells by Luo et
negatively affect the digestive enzymes including al. (2010). Experimental evidences showing the
trypsin, chymotrypsin and amylase (Singh 1988). hypocholesterolemic effect of stilbenes-containing
Compounds like phytolectins are heat-labile extracts from pigeonpea have also been furnished
hence get destroyed easily while cooking (Saxena by Luo et al. (2008). The wide ranging biological
et al., 2010a). Concerning its therapeutic uses, activities and medicinal properties of diverse
consumption of immature pigeonpea seeds has chemical compounds obtained from pigeonpea
been considered to have ameliorating effects in were reviewed recently by Pal et al. (2011).
7. Known interactions with other organisms in

managed and unmanaged ecosystems
7.1 Interactions in Unmanaged spaces in forests and creeping types climb on the
and Managed Ecosystems trees to get light. However, pigeonpea has not been
known to grow as naturalized population, hence
Pigeonpea prefers grassy habitats in tropical and
sub-tropical cold free zones with an optimum its interaction in the unmamaged ecosystem is
annual rain fall of 600-1000 mm. It can also uncertain. Stabilizing yield in pigeonpea is a major
grow in open areas in the forests, hilly slopes concern as its production is very much affected
and degraded lands under natural habitat. Wild by several biotic and abiotic factors (Varshney et
pigeonpea colonizes the drained, sunny and open al., 2013). Pigeonpea is grown as a rainfed crop

and precipitation affects this crop differentially in storage, bruchids are the most dangerous. The
different regions depending on the soil type, slope insect pests of pigeonpea are briefly described here:
of land and natural drainage. In the absence of
(i) Helicoverpa pod borer (Helicoverpa arm-
proper rains the crop may fail or its yield will be
igera): This insect pest
severely affected; excess rain is also very harmful is also known as gram
to this crop in the early growth stages (up to 60 pod borer (Fig 8). The
days). In the poorly drained fields heavy rain larvae damage the pods or
immediately after sowing may lead to heavy loss flower buds and feed on
of plant stand which subsequently will affect the leaves as well. The typical
yield. Waterlogging during initial growth stage, circular hole on pods can
low temperature during flowering stage, high be observed through which
temperature during pod formation and drought are the seeds are damaged by Fig. 8: Gram pod
impediments in realizing the productivity potential the pod borer. borer
of pigeonpea cultivars. Although pigeonpea has
(ii) Maruca pod borer (Maruca vitrata) : It is
deep roots, yield losses due to these stresses are
an insect pest causing
large and widespread, especially when they occur
huge damage especially to
during critical seedling and reproductive stages.
short duration pigeonpea
in different parts of the
7.2 Important Insect Pests, Nature country. Usually the larvae
of Damage and their control in of this insect web tender
Managed Ecosystems leaves along with buds,
flowers and immature Fig. 9: Maruca pod
7.2.1 Major Insect Pests
pods and feed on them, thus borer
A variety of insect species (over 200) affects causing substantial economic damage (Fig 9).
pigeonpea plant and seeds, and feeds on roots, (iii) Pod Fly (Melanagromyza obtusa): It is
shoots, flowers, and seeds. However, majority of considered to be one of the dreaded pests of
these insects are sporadic in their distribution and pigeonpea. The maggot of the insect feeds
do not cause economic damage and therefore, may on the developing grain
not be regarded as pests. In general, pigeonpea (Fig 10). The infested
can tolerate foliage damage during vegetative pods do not show
growth and need no chemical control. The any external evidence
damage caused to reproductive parts is difficult of damage until these
to compensate in short duration varieties, while maggots are fully grow
in long duration cultivars recovery from damage and larvae make holes
is slow and dependent on the plant type, soil in the pod walls. The
moisture and climatic conditions. maggots bore the grains
and make tunnels in Fig. 10: Maggot
The devastating insect pests which attack pigeonpea them. This hole provides feeding on
at the reproductive phase (flower buds, flowers, an emergence “window” developing grain
pods, and seeds) are pod borer and pod fly. During through which the adults exit the pod.

Damaged grains or seeds do not mature and penetrate the pod wall and suck the liquid
fungus can be seen on excreta inside pods. from developing seeds. Damaged seeds
The infested seeds or grains lose their viability become shriveled, and develop dark patches.
and do no germinate. Seeds spoiled by pod sucking bugs neither
(iv) Blister beetle (Zonabris pustulata): This pest germinate nor are acceptable as human food.
feeds voraciously on the flowers of pigeonpea (viii) Red spider mite (Tetranychus spp.): Spider
mites cause yellow or white spots on the
upper surface of the infested leaflets. Heavy
infestation results in bronzing of the leaves
followed by defoliation.
(ix) Eriophyid mite (Aceria cajani): The
eriphyid mite, A. cajani is the vector of the
pigeonpea sterility mosaic disease (SMD),
the most serious viral disease of this crop.
Plants infected with sterility mosaic disease
Fig. 11: Blister betle develop light green, chlorotic foliage. The
and greatly reduces the pod setting (Fig 11). early infection results in reproductively sterile
This insect can cause huge damage in case of plants.
favourable conditions i.e. high humidity and
mild temperature. It is a serious pest in short 7.2.2 Major Nematodes
duration varieties. Nematodes are widely distributed in most of the
(v) Pigeonpea plume moth (Exelastis atomosa pigeonpea growing regions in India. Pigeonpea is
Wals.): The larvae damage seeds as well as vulnerable to many plant parasitic nematodes viz.,
cause flowers, buds and pods to drop. Small Meloidogyne spp., Heterodera cajani, Helicotylenchus
spiny greenish brown caterpillar and pupae spp., Hoplolaimus spp., Rotylenchus spp. and
can be seen on the pods. It also enters into Tylenchorhynchus spp. Root Knot Nematode (M.
pod and feeds on developing grains resulting javanica) and pigeonpea cyst nematode (Heterodera
in reduced yields. cajani) are the most important parasites.
(vi) Pigeonpea blue butterfly (Euchrysops cnejus, The lower larval population of H. arimgera,
Lampides boeticus and Catochrysops strabo): E. atomosa and L. boeticus as well as their damage were
The larvae feed on flowers, seeds and pods of recorded, when crop was treated with indoxacarb
pigeonpea. Specific control for these insects is (0.007%), which was at par with spinosad (0.009%),
rarely required but the general management fenvalerate (0.01%) and monocroptophos (0.36%)
recommendations for H. armigera may be (Ghetiya and Mehta, 2011).
used here.
(vii) Pigeonpea pod sucking bug (Clavigralla 7.2.3 Integrated Pest Management Practices in
gibbosa): The adults and nymphs insect, Pigeonpea
of this use their piercing mouthparts to 1. Deep ploughing during summer to expose the

hibernating pupae of pod borers to adverse seeds can also transmit the disease. Symtoms like
weather conditions and natural enemies. drooping and subsequent drying of the plants
2. Mixing of sorghum/maize seeds (250-500 g/ can be observed in field conditions (Fig 12). The
ha) to function as live bird perches. These stem of infected plant when cut vertically, show
plants also help in conserving natural enemies. black lines which indicate infection of Fusarium
wilt.
3. Installation of pheromone traps @ 40/ha and
replace helilure at an interval of 21 days as (ii ) Sterility mosaic disease (Pigeonpea sterility
per need. mosaic virus)
4. Application of HaNPV @ 250 LE/ha protects
the crop from H. armigera. SMD causes stunting, small yellowish green
leaves, bushy habit and complete cessation of
5. Spraying of monocrotophos 36 SL @ 500 ml/
growth of reproductive structures. The degree
ha is effective against pod fly and pod borers.
of sterility due to the suppression of flowers and
Since pigeonpea is ravaged by an array of pest fruits may vary in different pigeonpea cultivars
species, the use of specific bio-pesticides that reduce (Sharma et al., 2015). The disease has now spread
the population of one or two key insects will not to different pigeonpea growing areas particularly
control the pest problem completely. Therefore, in Bihar, Karnataka, Tamil Nadu, Gujarat and
the role of synthetic insecticides in suppressing Maharashtra. Management of the disease through
pest complex infesting pigeonpea will continue cultural practices is not well
to play an important role. However, integrated studied.
pest management involving tolerant varieties,
(iii) Phytophthora blight
monitoring through pheromone traps, need based
(Phytophthora drechsleri
use of insectides and crop rotations and other
Tucker f. sp. Cajani)
agronomic manipulations need to be adopted to
control damages of pests. In addition to fusarium wilt
and SMD, phytophthora Fig. 13: Foliar disease affecting stem
7.3 Important Diseases, Causal blight is another important
agents and their Control in disease of pigeonpea (Pande et al. 2011). It is a
Managed Ecosystems foliar disease affecting both leaves and stems (Fig
13). The disease can appear as soon as the pigeonpea
(i) Fusarium wilt
seedlings emerge and can go unnoticed as the small
(Fusarium udum Butler)
germinated seedlings collapse on the ground with
Fusarium wilt is the most “damping - off ” type of symptoms. The disease in
important and wide the field is usually recognised when the seedlings are
spread fungal disease in about two weeks old. Water soaked lesions appear
all pigeonpea growing on the leaves or breaking of stem with broken upper
areas (Pande et al., 2013). part of the plant still attached to the basal portion of
The primary source of the stem. With close observation, brown, shrunken
Fig. 12: Drooping of plant innoculum is soil. Infected lesions can be seen on the above ground part of the

stem even before the breaking of the stem occurs. lowlying areas of the field and in waterpaths as it is
In the grown-up plants, development of cankerous promoted by higher humidity. The mortality can be
out growths (gall) on the edges of the stem lesions reduced by sowing pigeonpea on ridges permitting
is common. The disease usually appears in the less splashes of rains as well as better drainage.
8. agronomic practices
8.1 Climate onset of the monsoon. Early sowing of short

duration varieties in the first fortnight of June
Various cultivars of pigeonpea are grown from
with irrigation or pre-monsoon rains gives higher
sea level up to 3,000 m altitude (Van Den Beldt,
seed yield. Delayed sowings result in progressive
1988). Reddy and Virmani (1981) reported that
reductions in yield due to early flowering and slow
pigeonpea can be grown between 14°N and 28°N growth. Late sowings of short duration varieties
latitude, with a temperature ranging from 26° to may result in poor pod and grain development,
30°C in the rainy season (June - October) and due to onset of cold season at the pod-filling
17° - 22°C in the post rainy (November - March) stage. Sowing in first fortnight of June also ensures
season. Pigeonpea is very sensitive to low radiation availability of field for post rainy season crops by
at flowering and pod development. Therefore, the end of November. Therefore, sowing should
flowering during the monsoon and cloudy weather not be delayed beyond June for early maturing
lead to flower/bud drop and poor pod formation. varieties. For medium duration varieties of
Pigeonpea is comparatively tolerant to drought central and southern zones, sowing at the
and high temperatures as compared to other pulse onset of rains (15th June - 15th July) ensures
crops. Although the plant can survive in very dry sufficient vegetative growth before flowering.
conditions, it has been observed that seed yield The long duration varieties should be planted
is minimal under these conditions. Pigeonpea is between second and last week of July in the
not suitable for heavy rainfall areas unless there is North East plains. Late sowings produce less
proper drainage system. Pigeonpea can survive at vegetative growth and may expose the crop to
low temperature to a certain degree but it is highly terminal drought and heat stress. September is
sensitive to frost damage which causes burn injury the optimum time for post-rainy season sowing
and leads to heavy defoliation followed by drastic in peninsular India. Delay in sowing affects the
delay in reproductive phase resulting in negligible vegetative growth and exposes the crop to high
pod set. temperature during the reproductive phase. The
recommended crop management practices may
8.2 Sowing Season be followed to raise a good crop.
Pigeonpea varieties are manily categorised in
8.3 Seed Production
three maturity groups (short, medium and long
duration), and planting is usually done with the Seed is the most important input of the crop

production. Seed may be defined as the propagating standardized and well organized conditions.
material comprising any part of the plant that During seed production strict attention is given
grows into a new plant and makes a link between to maintain the genetic purity and other qualities
previous generation and present generation. Seed of the seed. Genetic purity of seed is maintained
production is the art of producing genetically pure by regular rouging of off-type plants. For the
plant propagules in large quantity with the aid of purpose of identifying off-types, the nodal person
science and technology. It is important that seed involved in seed production must possess the list
of a new and superior variety should be multiplied of diagnostic traits of the variety. Availability of
and made available in adequate quantities as soon quality seed of improved varieties has been a major
as possible so as to benefit the farming community. constraint. Package of practices for raising a seed
Seed of released varieties must be maintained and crop remains same for pure line or hybrid seed
kept ready for pushing them into the commercial production. The crop management practices for
seed chain. Seed production is carried out under raising a good crop are given in Table 5.
Table 5. Crop management practices for raising a good crop
Crop management
Seed treatment Seed treatment with carbendazim 50 WP) 2g + thiride 75 WP) 2g +
Metalaxyl-M 45.3%) 2g per Kg seed
Soil treatment Trichoderma viride @ 2.5 kg/ha+ Farm Yard Mannure + neem cake @ 5 q/ha
Carbofuran @ 1Kg ai/ha
Fertilizers Basal application 20 kg N + 40-60 kg P2O5+ 20Kg K2O + 10 Kg ZnSO4
in 1 ha area
Weeds Pre-emergence application of pendimethalin @ 1.5 kg/ha
Two hand weedings at 30 and 60 DAS
Disease management 1. Deep ploughing and exposing field to sun during hot summer months
2. Field sanitation, removal of all stubbles and plant remains
3. Ridge sowing
4. Destruction of off-season volunteer plants from the vicinity of seed plot
Sterility mosaic disease Preventive measure of acaricide Dicofol 18.5% EC @ 2.5 ml per liter water
Fusarium wilt See treatment using fungicide Carbendazim @2.5 g/kg of
seed/Trichoderma harzianum @6-10g/kg
Alternaria blight In the event of disease, foliar spray of mancozeb @ 0.2%
Phytophthora blight As a preventive measure: foliar spraying of Metalaxyl-M 45.3 % @ 3g/l at 30
and 45 days after sowing.
Pod borer and pod fly control At podding stage, 2-3 foliar sprays of Spinosad 45SC @ 0.4-0.5 ml/liter at an
interval of 15 days

9. Breeding objectives
9.1 Milestones in Breeding identification of genetic male sterility offered

new avenues for expoiting heterosis leading to
In the pre independence time all cultivated
the initiation of ICAR sponsored programme
pigeonpea genotypes were landraces or local types
on “Promotion of Hybrids in Selected Crops” in
which were either long duration type (>180 days)
1989. Heterosis breeding was tried for breaking
in the Northern plains or medium late in the
the yield barrier through exploitation of hybrid
central and southern zone. With the creation of
vigour. Initially genetic male sterility (GMS) was
AICPIP, pigeonpea breeding efforts were initiated
utilised for the development of hybrids and, as a
simultaneously at 31 locations in different
result an early maturing hybrid ICPH 8, which
agroclimatic zones of the country for developing
showed more than 40 % superiority over the best
early, medium and long duration varieties suitable
check UPAS 120, was released for commercial
for different zones (Ramanujam and Singh, 1981).
cultivation (Saxena et al., 1992). Subsequently,
Multilocation testing of genotypes for yield and
5 more GMS based hybrids were released for
biotic stresses paved the way for the identification
general cultivation. Though ICPH 8 and other
of more stable varieties. Varieties were bred either
hybrids had substantial yield advantage, they could
through hybridization followed by pure line
selection or by selecting desirable plants from the not survive due to difficulties in seed production
heterogeneous germplasm followed by pure line associted with GMS system that led to very high
selection. During this time a need for reducing cost of hybrid seed (Saxena et al., 2010b). In order
the crop duration was felt. UPAS120 was the to overcome the lacuna of GMS, cytoplasmic
first high yielding short duration (120-140 days) genetic male sterility (CGMS) system was adopted
crop released in 1976. ICRISAT has played an for the production of hybrid seed. The hybrid
important role in the introduction of medium programme was further strengthened through the
and short duration varieties for the Central and National Agricultural Technology Project (NATP)
Southern zone. About 40% of the varieties bred in 1998 and the Integrated Scheme on Oilseeds,
so far have evolved through selections made Pulses, Oilpalm and Maize (ISOPOM) in 2005.
from landraces or heterogeneous population or In 2004, the first CMS based medium duration
spontaneous mutations. Vishakha 1 (TT 6) was the hybrid GTH 1 was released in India (Varshney
first variety developed through mutation breeding et al., 2010). Subsequently, using A4 cytoplam
by BARC. Subsequently, three more varieties were a medium duration hybrid RVICPH 2671 was
developed through mutation breeding. Inspite of released in Madhya Pradesh.
long and continued breeding efforts, the average
productivity of pigeonpea has not increased 9.3 Major Traits of Interest and
significantly in the last five decades. Priorities in Breeding
The important agronomic and yield attributing
9.2 Heterosis Breeding in
traits of pigeonpea for enhancing productivity are:
Pigeonpea days to 50% flowering, days to 75 % maturity, plant
Existence of high natural out crossing and height, plant type, branching pattern, number of

pods per plant, number of seeds per pod, pod BSMR 736, BSMR 853, ICP 7035 also show
length, number of primary branches, number of notable levels of resistance against SMD. Recently,
secondary branches, branching length, 100 seed screening of pigeonpea core collection comprising
weight, seed yield per plant and seed yield per plot 146 accessions have revealed a set of 24 accessions
(Saxena, 2008). Tolerance against biotic and abiotic that have promising levels of resistance to SMD.
stresses is an important parameter for selection. More important, five accessions exhibited marked
The objectives of the breeding programme resistance not only to SMD but also against
depend on the local needs of the cultivators, Fusarium wilt (Sharma et al., 2012). In case of
prevailing cropping system, climatic condition Phytophthora blight, only a limited number of
and constraints of production. Though breeding promising lines like KPBR 80 and ICP 9252 have
objectives vary in different agroclimatic zones as per been reported so far, which show resistance against
the local needs, varieties are generally bred keeping specific races (Saxena, 2005).
in view appropriate maturity and enhanced harvest
index through manipulating plant architecture 9.4 Advancements and Challenges
for different cropping situations. Attention is also
Cleistogamy favours self pollination in pigeonpea
given to induce photo insensitivity, enhancing
but it exhibits a range of natural out crossing
yield and ensure stability of the released varieties
rate due to insect aided pollination (Saxena and
by incorporating resistance against Fusarium wilt,
Kumar, 2010). There are no reports of inbreeding
SMD and Phytopthora blight and tolerance against
depression and this crop exhibits commercially
abiotic stresses like drought, heat and cold.
exploitable level of heterosis which is why major
Identification of potential donors and incorporation breeding methodologies so far adopted are based on
of disease resistance into newly developed cultivars exploiting additive genetic variance. Considerable
remains the most sustainable, ecomically attractive advancement has been made in extracting F1
and ecofriendly strategy to reduce yield losses heterosis through the development of hybrids
in crop plants. With the aim of identifing the in short and medium duration group. Pure line
Fusarium wilt resistant sources, multi-location breeding has been the main strategy for genetic
and multi-year screening of genotypes led to enhancement, but population improvement has
the identification of promising wilt resistance also been tried in pigeonpea (Khan, 1973; Onim,
genotypes in pigeonpea like IPA 16 F, IPA 8 F, 1981b). It was suggested that out-crossing potential
IPA 9 F and IPA 12 F (Singh et al., 2011). Several of pigeonpea should be utilized in the formation
varieties have been developed in pigeonpea like of random mating composites which will serve
ICP 8863, ICPL 87119, BDN 1, BDN 2, BSMR as the dynamic reservoir of variability and can
736, IPA 203 which are known to exhibit resistance also be used as base populations for studies in
against Fusarium wilt (Choudhary et al., 2013). natural selection, mass selection and recurrent
Similarly, Kulkarni et al. (2003) reported some selection. Stratified mass selection and mass
C. scarabaeoides accessions, which were found to selection with progeny testing were used for yield
be resistant to pigeonpea sterility mosaic virus. improvement but very little response was observed
Among the pigeonpea lines commercially available (Onim, 1981b). Pigeonpea has shown substantial
for cultivation, popular varieties like ICPL 87119, amount of non-additive genetic variance and

hybrid vigour for yield. The discovery of stable in pigeonpea. Recently, CMS based (three
genetic male sterility coupled with its out crossing line system; ‘A’, ‘B’ and ‘R’) hybrids have been
nature, has opened the possibility of commercial released for cultivation. The ‘B’, ‘R’ and the
utilization of the heterosis in pigeonpea (Saxena released varieties of pigeonpea are purelines and,
et al., 2010b). The hybrid technology, based on hence, their maintenance and seed production
cytoplasmic nuclear male-sterility system, has given techniques are same. However, specialized
an opportunity for achieving the long-cherished techniques are employed to maintain ‘A’ line
goal of breaking yield barrier in pigeonpea (Saxena (CMS line) and to produce F1 hybrid seed.
and Nadarajan, 2010). The short and medium- Several biotic factors severely constrain the
duration types and disease resistant cultivars have productivity of pigeonpea (Varshney et al.,
made a significant impact on increasing area under 2010, 2013). The losses caused by diseases can
pigeonpea cultivation. be effectively controlled through host-plant
resistance breeding (Sharma et al., 2012). The
In the national agricultural research system major challenges ahead are to enhance genetic
(NARS), genetic improvement in pigeonpea yield potential and manage damages are being
has been brought primarily through pure caused by the gram pod borer and other insects
line breeding. Population improvement and some of the diseases where high level of
programmes, tried elsewhere, have not been genetic resistance is not available within the
successful. After long continued efforts of pigeonpea germplasm (cultivated and wild
heterosis breeding, hybrids are now a reality ones).
10. Biotechnological interventions in

pigeonpea
10.1 Transgenic Approaches for gene transfer, various methods like Agrobacterium-
Genetic Improvement mediated, biolistic and electroporation have been
developed to facilitate foreign DNA transfer in
Transgenic technology offers a means to plants (Eapen, 2008).
introgressing traits that are not amenable to transfer
using conventional breeding techniques or adequate Among the methods mentioned above,
variability is not available in the exploitable Agroacterium mediated particularly using
gene pool. However, poor transformation and A. tumefaciens strain ‘LBA4404’ has been widely
regeneration capacities along with differential employed in pigeonpea (Geetha et al., 1999,
response of genotypes to regeneration protocols Surekha et al., 2007). In contrast, Dayal et al.
offer the major impediments to development (2003) developed a reliable regeneration protocol
of transgenic pigeonpea (Geetha et al. 1999; using leaves as explants and micro-particle
Lawrence and Koundal, 2001; Rao et al., 2008; bombardment method for DNA transfer. Thu
Eapen, 2008). Differential reactions to various A. et al. (2003) achieved transformation with both
tumefaciens strains were also elucidated (Surekha et A. tumefaciens mediated and micro-particle
al., 2007; Chandra and Pental, 2003). Concerning bombardment-driven gene transfer. Lawrence and

Koundal (2001) transformed pigeonpea plant with mapping populations segregating for agronomic
CaMV 35S promoter-led cowpea protease inhibitor traits including SMD and FW resistance and
gene. Similarly, Kumar et al. (2004) used hpt and fertility restoration were developed (Varshney et al.,
rice chitinase genes in pigeonpea transformation. 2010). To this end, Bohra et al. (2011) reported
To generate pest-resistant pigeonpea. Sharma et al. high throughput development of SSR markers via
(2006) demonstrated the successful integration of mining bacterial artificial chromosome (BAC)-end
the Bt cry1Ab gene. Successful regeneration has sequences (Bohra et al., 2011). The 3,072 BES-SSR
been obtained in pigeonpea using both ways viz. markers were obtained, which were subsequently
organogenesis and somatic embryogenesis (Patel et used in various genomics application like genetic
al., 1994; George and Eapen, 1994). In addition, linkage mapping and QTL analysis.
direct regeneration as well as callus induction have
also been reported in pigeonpea (Lawrence and The first SSR based genetic map was reported
Koundal, 2001; Kumar et al., 1983; Chandra and for an inter-specific mapping population (Bohra
Pental, 2003). Broad range of explants including et al., 2011). Later, a series of SSR based genetic
cotyledonary nodes, shoot apices, decapitated maps were constructed for different intra-specific
embryonic axis, leaf etc. has been used for in vitro F2 populations. A number of SSR markers were
regeneration of transformed plants in pigeonpea successfully mapped ranging from 59 (ICPB
(Eapen, 2008; Chandra and Pental, 2003). As an 2049 × ICPL 99050) to 140 (ICPA 2043 × ICPR
alternative to escape the labour intensive and low 3467) spanning 586 and 881.6 cM, respectively.
throughput in vitro regeneration, Rao et al. (2008) In parallel, these mapping populations were used
developed in planta transformation system for for discovery of the gene(s)/QTL(s) governing the
pigeonpea. In this investigation, a total of 48 plants traits of interest (Bohra et al., 2012). These QTLs
generated PCR products for both gus (uid A) and explained substantial amount of the phenotypic
npt II genes, and the integration and transmission variation for the trait under investigation. QTLs
were confirmed by Southern blot analysis. restoring fertility accounted to 24% of the variation
However, even after using all above mentioned from an F2 population ICPA 2043 × ICPR 3467.
techniques, there is no transgenic cultivar available Similarly, six QTLs linked with the SMD resistance
as of now for cultivation possessing abiotic or biotic were also discovered from two different populations
stress resistance in pigeonpea. viz. ICP 8863 × ICPL 20097 and TTB 7 × ICP
7035 (Gnanesh et al., 2011). In addition to SSRs,
10.2 Genomics and Molecular other markers like random amplified polymorphic
Breeding in Pigeonpea DNA (RAPD) and amplified fragment length
Availability of genomic resources like robust polymorphism (AFLP) were also found to be
markers/quantitative trait loci (QTLs) sets a associated with Fusarium wilt, Plant type and
prerequisite for undertaking molecular breeding. In SMD resistance using bulked segregants analysis
pigeonpea, handful of SSRs was available in public (BSA) (Dhanasekar et al., 2010; Ganapathy et al.,
domain till 2010. However, significant research 2009; Kotresh et al., 2006). Recently, Kumawat
investments have been made during the last five et al. (2012) detected QTLs for plant type and
years and due to which tremendous genomic earliness in an F2 population viz Pusa Dwarf ×
resources have been generated. As a result, several HDM04-1.

A popular pigeonpea cultivar ‘Asha (ICPL 87119)’ pigeonpea genotypes resulted in identification of
was chosen for sequencing the whole genome by functionally-relevant SNP markers. As a result,
using NGS platforms including 454 GS-FLX 28,104 novel SNPs were added to the marker
(Singh et al., 2011) and Hiseq 2000 Sequencing repertoire of pigeonpea. The genotypes involved in
Systems (Varshney et al., 2012). Approximately, 72 SNP identification represented parents of different
% (606 Mb) and 61% (~511 Mb) of the pigeonpea mapping population that segregate for various
genome was successfully assembled in the two important traits (Varshney et al., 2012). Cost
separate sequencing attempts by Varshney et al. effective 116 KASPar assays for pigeonpea were
(2012) and Singh et al., (2011), respectively. A also developed recently to study the level of genetic
total of 48,680 protein coding genes were predicted variability and for other applications (Saxena et
in the genome and 51.67% of the genome was al., 2014). Likewise, after experimental validation,
represented by repetitive elements (Varshney et Singh et al. (2011) provided a set of SSR markers
al., 2012). Similar number of protein coding genes comprising 437 markers and designated these
(47,004) was reported by Singh et al. (2011). The markers as hypervariable ‘Arhar’ simple sequence
number of protein coding genes was similar to repeat (HASSR) markers. Certainly, all these
those observed in the genomes of other related predictive molecular markers will be of great help
legumes species as well as rice and Arabidopsis. to pigeonpea breeders for accelerating the progress
Furthermore, in silico mining of the whole genome of crop improvement. In addition to pigeonpea
sequence permitted access to large-scale DNA genome sequencing, mitochondrial genomes of four
markers especially SSRs and SNPs (Varshney et Cajanus genotypes : the CMS line ICPA 2039,
al., 2012). A total of 309,052 SSRs were detected its cognate maintainer line ICPB 2039, the hybrid
across the genome, of which primer pairs were line ICPH 2433 and the wild relative ICPW 29
successfully designed for 23,410 SSRs. In parallel, (accession from C. cajanifolius), source of A4
analysis of the transcript reads from 12 different cytoplasm were also sequenced (Tuteja et al., 2013).

11. References
Andersson MS, de Vicente MC (2010) Gene Flow Between Crops And Their Wild Relatives. 564 pages. Johns
Hopkins University Press, Baltimore, MD.
Bahadur B, Ra o MM, Rao KL (1981) Studies on dimorphic stamens and pollen (SEM) and its possible role
inpollination biology of Cajanus cajan Millsp. Indian Journal of Botany 4: 122-129.
Bhatia GK, Gupta SC, Green JM, Sharma D (1981) Estimates of natural cross-pollination in Cajanus cajan (L.)
Millsp. Several experimental approaches International Workshop on Pigeonpeas. pp 129–136, vol 2, 15–19 Dec
1980. ICRISAT, Patancheru, India.
Bhatia VS, Singh P, Wani SP, Kesava Rao AVR, Srinivas K (2006) Yield Gap Analysis of Soybean, Groundnut,
Pigeonpea and Chickpea in India Using Simulation Modeling. Global Theme on Agro-ecosystems. Report no.
31. 156 pp, Andhra Pradesh, India: International Crops Research Institute for the Semi-Arid Tropics (ICRISAT)
Patancheru, India.
Bisen SS, Sheldrake AR (1981) The anatomy of the pigeonpea. Research Bulletin 5, International Crops Research
Institute for the Semi-Arid Tropics, Patancheru, India.
Bohra A, Mallikarjuna N, Saxena KB, Upadhyaya H, Vales I, Varshney R (2010) Harnessing the Potential of Crop
Wild Relatives Through Genomics Tools for Pigeonpea Improvement. Journal of Plant Biology 37: 83-98.
Bohra A, Dubey A, Saxena RK, Penmetsa RV, Poornima KN, Kumar N, Farmer AD, Srivani G, Upadhyaya HD,
Gothalwal R, Ramesh R, Singh D, Saxena KB, Kavi Kishor PB, Singh NK, Town CD, May GD, Cook DR, Varshney
RK (2011) Analysis of BAC-end sequences (BESs) and development of BES-SSR markers for genetic mapping and
hybrid purity assessment in pigeonpea. BMC Plant Biology 11:56
Bohra A, Saxena RK, Gnanesh BN, Saxena KB, Byregowda M, Rathore A, Kavi Kishor PB, Cook DR, Varshney
RK (2012) An intra-specific consensus genetic map of pigeonpea [Cajanus cajan (L.) Millspaugh] derived from six
mapping populations. Theoretical and Applied Genetics 125:1325-1338.
Chandra A, Pental D (2003) Regeneration and genetic transformation of grain legumes: An overview. Current
Science 84:381-387
Chikowo R, Mapfumo P, Nyamugafata P, Giller KE (2004) Woody legume fallow productivity, biological N2-fixation
and residual benefits to two successive maize crops in Zimbabwe. Plant and Soil 262: 303–315
Choudhary AK, Kumar S, Patil BS, Bhat JS, Sharma M, Kemal S, Ontagodi TP, Datta S, Patil P, Chaturvedi SK,
Sultana R, Hegde VS, Choudhary S, Kamannavar PY, Vijayakumar AG (2013) Narrowing yield gaps through
genetic improvement for Fusarium wilt resistance in three pulse crops of the semi-arid tropics. SABRAO Journal
of Breeding and Genetics 45: 341-370.
Dayal S, Lavanya M, Devi P, Sharma KK (2003) An efficient protocol for shoot regeneration and genetic
transformation of pigeonpea [Cajanus cajan (L.) Millsp.] using leaf explants. Plant Cell Reports 21:1072–1079

De DN (1974) Pigeonpea. In: Evolutionary Studies in World Crops, Diversity and Change in the Indian Subcontinent
(Hutchinson J Ed.), pp. 79-87, Cambridge University Press, London.
Dhanasekar P, Dhumal KH, Reddy KS (2010) Identification of RAPD marker linked to plant type gene in pigeonpea.
Indian Journal of Biotechnology 9:58–63.
Dhanju MS, Gill BS (1985) Intergeneric hybridization between Cajanus cajan and Atylosia platycarpa. Annals of
Biology 1:229–231.
Dundas IS (1984) Cytogenetic investigations involving pigeonpea (Cajanus cajan (L.) Millsp.) and some Australina
Atylosia specie. Ph.D. thesis, Department of Agriculture, University of Queensland, Australia.
Dundas IS (1990) Pigeonpea: cytology and cytogenetics – perspectives and prospects. In: The Pigeonpea (Nene YL,
Hall SD, Sheila VK eds.), pp. 117—136. CAB International, Wallington, UK.
Dutta PC, Deb A (1970) Floral biology of Cajanus cajan (Linn.) Millsp. var. Bicolor D.C. (Papilionaceae). Bulletin
of the Botanical Society of Bengal 24: 135-145.
Eapen E (2008) Advances in development of transgenic pulse crops. Biotechnology Advances 26:162–168
FAOSTAT (2013). Food and Agriculture Organization of the United Nations. FAOSTAT. http://faostat3.fao.org/
home/index.html
Ganapathy KN, Byregowda M, Venkatesha SC, Rama Chandra R, Gnanesh BN, Girish G (2009) Identification
of AFLP markers linked to sterility mosaic disease in pigeonpea Cajanus cajan (L.) Millsp. International Journal
of Integrative Biology 7: 145-149.
Ganeshan S (2008) Traditional oral care medicinal plants survey of Tamil nadu. Natural Product Radiance 7:166–72.
Geetha N, Venkatachalam P, Lakshmi Sita G (1999) Agrobacterium-mediated genetic transformation of pigeonpea

(Cajanus cajan L.) and development of transgenic plants via direct organogenesis. Plant Biotechnol 16:213–218
George L, S Eapen (1994) Organogenesis and embryogenesis from diverse explants in pigeonpea (Cajanus cajan
L.). Plant Cell Reporter 13: 417–420.
Gill LS, Hussaini SWH (1986) Cytological observations in Leguminosae from Southern Negeria, Willdenowia,
15, 521.
Gnanesh BN, Bohra A, Sharma M, Byregowda M, Pande S, Wesley V, Saxena RK, Saxena KB, KaviKishor PB,
Varshney RK (2011) Genetic mapping and quantitative trait locus analysis of resistance to sterility mosaic disease
in pigeonpea [Cajanus cajan (L.) Millsp.]. Field Crops Research 123:53–61
Gooding HJ (1962) The agronomic aspects of pigeonpeas. Field Crop Abstracts 15:1–5.
Gowda CLL, Saxena KB, Srivastava RK, Upadhyaya HD, Silim SN (2011) Pigeonpea: From an orphan to leader
in food legumes. In Biodiversity in Agriculture: Domestication, Evolution and Sustainability (Gepts P, Bettinger
RL, Brush SB, Famula TR, McGuire PE, Qualset CO and Damania AB eds.). Pp 361-373. Cambridge University
Press. University of California, Davis, USA.

Greilhuber J, Obermayer R (1998) Genome size variation in Cajanus cajan (Fabaceae): a reconsideration. Plant
Systematics and Evolution 212:135–141
Harlan JR, de Wet JMJ (1971) Towards a rational classification of cultivated plants. Taxon 20: 509–517.
Howard A, Howard GLC, Khan LN (1919) Studies in the pollination of Indian Crops.Memoirs of the Department
of Agriculture in India (Botanical Senes) 10: 201-207.
Jayaprakash P, Sarla N (2001) Development of an improved medium for germination of Cajanus cajan (L.) Millsp.
pollen in vitro. Journal of Experimental Botany 52: 851–855
Jha SS, Ohri D (1996) Phylogenetic relationships of Cajanus cajan (L.) Millsp. (Pigeonpea) and its wild relatives.
Genetic Resources and Crop Evolution 43:275–281
Kassa MT, Penmetsa RV, Carrasquilla-Garcia N, Sarma BK, Datta S, Upadhyaya HD, et al. (2012) Genetic patterns
of domestication in pigeonpea (Cajanus cajan (L.) Millsp.) and wild Cajanus relatives. PLoS One 7:e39563.
KhanTN (1973) A new approach to the breeding of pigeon pea (Cajanus cajan Millsp.): formation of composites.
Euphytica 22:373-377.
Kollipara KP, Singh L, Hymowitz T (1994) Genetic variationof trypsin and chymotrypsin inhibitors in pigeonpea,
Cajanus cajan (L.) Millsp, and its wild relatives. Theoretical and Applied Genetics 88:986–993.
Kotresh H, Fakrudin B, Punnuri S, Rajkumar B, Thudi M, Paramesh H, Lohithswa H, Kuruvinashetti M (2006)

Identification of two RAPD markers genetically linked to a recessive allele of a Fusarium wilt resistance gene in
pigeonpea (Cajanus cajan (L.) Millsp.). Euphytica 149:113–120.
Krauss FG (1936) Pigeon peas as human food and as a feed for livestock.Univ.Hawaii Coop. Ext.Service. Agricultural
Notes AN-125, 4 pp.
Krishna TG, Reddy LJ (1982) Species affinities between Cajanus cajan and some Atylosia species based on esterase
isozyme. Euphytica 31:709–713
Krishnaswamy N, Ayyangar GNR (1935) Chromosome number in Cajanus indicus Spreng. Current Science 3: 614.
Kulkarni NK, Reddy AS, Kumar PL, Vijayanarasimha J, Rangaswamy KT, Muniyappa V, Reddy LJ, Saxena KB,
Jones AT, Reddy DVR (2003) Broad based resistance to pigeonpea sterility mosaic disease in accessions of Cajanus
scarbaeoides (L.) Benth. Indian Journal of Plant Protection 31: 6-11.
Kumar AS, Reddy TP, Reddy GM (1983) Plantlet regeneration from different callus cultures of pigeonpea (Cajanus
cajan L.). Plant Science Letters 32: 271–278.
Kumar LSS, Thombre MV, D’Cruz R (1985) Cytological studies of an inter-generic hybrid of Cajanus cajan (L)
Millsp. and Alytosia lineata. Proceedings of the National Academy of Sciences, India, Section B 47: 252.
Kumar PS (1985) Crossability, genome relationships and inheritance studies in intergeneric hybrid of pigeonpea.
Ph.D.thesis, University of Hyderabad, India.

Kumar, SM, Kumar K, Sharma KK, Devi P (2004) Genetic transformation of pigeon pea with rice chitinase gene.
Plant Breeding 123: 485–489
Kumawat G, Raje RS, Bhutani S, Pal JK, Mithra SVCR, Kishor Gaikwad K, Sharma TR, Singh NK (2012)
Molecular mapping of QTLs for plant type and earliness traits in pigeonpea (Cajanus cajan L. Millsp.). BMC
Genetics 13:84.
Lakshmi MP, Senthilkumar P, Parani M, Jithesh MN, Parida AK (2000) PCR-RFLP analysis of chloroplast gene
regions in Cajanus (Leguminosae) and allied genera. Euphytica 116: 243–250.
Lans C (2007) Comparison of plants used for skin and stomach problems in Trinidad and Tobago with Asian
Ethnomedicine. Journal of Ethnobiology and Ethnomedicine 33:22–8
Lawrence PK, Koundal KR (2001) Agrobacterium tumefaciens-mediated transformation of pigeonpea (Cajanus cajan
L. Millsp.) and molecular analysis of regenerated plants. Current Science 80:1428–1432.
Luo QF, Sun L, Si JY, Chen DH (2008) Hypochol-esterolemic effect stilbenes containing extract fraction Cajanus
cajan L diet induced hypercholesterolemia mice. Phytomedicine 15:932–939.
Luo M, Liu X, Zu Y, Fu Y, Zhang S, Yao L, Efferth T (2010) Cajanol, a novel anticancer agent from pigeonpea
[Cajanus cajan (L.) Millsp.] roots, induces apoptosis in human breast cancer cells through a ROS-mediated
mitochondrial pathway. Chemico-Biological Interactions 188:151–160
Mafongoya PL, Bationo A, Kihara J, Waswa BS (2006) Appropriate technologies to replenish soil fertility in southern
Africa. Nutrient Cycling in Agroecosystems 76:137–151.
Mallikarjuna N and Moss JP (1995) Production of hybrids between Cajanus platycarpus and Cajanus cajan. Euphytica
83: 43–46.
Mallikarjuna N (1998) Ovule culture to rescue aborting embryos from pigeonpea. Indian Journal of Experimental
Biology 36:225–228.
Mallikarjuna N and Saxena KB (2002) Production of hybrids between Cajanus acutifolius and C. cajan. Euphytica
124:107–110.
Mallikarjuna N and Saxena KB (2005) A new cytoplasmic male sterility system derived from cultivated pigeonpea
cytoplasm. Euphytica 142:143–148.
Mallikarjuna N, Deepak J, Prabhakar R (2006) Introgression of Cajanus platycarpus genome into cultivated pigeonpea,
C. cajan. Euphytica 149:161–167.
Mallikarjuna N (2007) Production of fertile progeny from interspecific incompatible cross Cajanus platycarpus ×
C. cajan. Journal of SAT Agricultural Research 3: 1-3.
Mallikarjuna N, Saxena KB, Jadhav DR (2011) Cajanus. In: Wild Crop Relatives: Genomic and Breeding Resources,
Legume Crops and Forages (C. Kole ed.), Springer-Verlag Berlin Heidelberg.

Mallikarjuna N, Saxena KB, Lakshmi J, Varshney R, Srikanth S, Jadhav D (2012) Differences between Cajanus
cajan (L.) Millspaugh and C. cajanifolius (Haines) van der Maesen, the progenitor species of pigeonpea. Genetic
Resources and Crop Evolution 59: 411-417.
Mazur WM, Duke JA, Kristiina W, Rasku S, Adlercreutz H (1998) Isoflavonoids and lignans in legumes: Nutritional
and health aspects in humans. Nutritional Biochemistry 9:193-200.
Nadimpalli, RG, Jarret JL, Pathak SC, Kochert G (1993) Phylogenetic relationships of pigeonpea (Cajanus cajan)
based on nuclear restriction fragment length polymorphism. Genome 36:216–223.
Nag YK, Sharma RN (2012) Genetic diversity and path coefficient analysis in Pigeonpea [Cajanus cajan(L.) Millsp.]
germplasm accessions of Bastar origin. Electronic Journal of Plant Breeding 3: 818-824.
Naithani ST (1941) Cytological studies on Indian pulses. Part I: The somatic chromosomes and the pro chromosomes
of Cajanus. Proceedings of the National Academy of Sciences, India 11:67–73.
Natarajan M, Willey RW (1980) Sorghum-pigeonpea inter cropping and the effect of plant population density on
growth and yield. Journal of Agricultural Science (UK) 95: 51-58.
Nwokolo E (1987) Nutritional evaluation of pigeon pea meal. Plant Foods for Human Nutrition 37:283-90
Odeny DA (2007a) The potential of pigeonpea (Cajanus cajan (L.) Millsp.) in Africa. Natural Resources Forum
31: 297–305.
Odeny DA, Jayashree B, Ferguson M, Hoisington D, Crouch J, Gebhardt C (2007b) Development, characterization
and utilization of microsatellite markers in pigeonpea. Plant Breeding 126:130–136.
Ohri D, Jha SS, Kumar S (1994) Variability in nuclear DNA content within pigeonpea. Plant Systematics and
Evolution 189:211–216
Ohri D, Singh SP (2002) Karyotypic and genome size variation in Cajanus cajan (L.) Millsp. (Pigeonpea) and some
wild relatives. Genetic Resources and Crop Evolution 49:1–10
Oke DG (2014) Proximate and Phytochemical Analysis of Cajanus Cajan (Pigeon Pea) Leaves. Chemical Science
Transactions 3: 1172-1178
Onim JFG (1981a) Pigeonpea improvement in Kenya. In:International Workshop on Pigeonpeas, vol. 1, 15–19
December1980, International Crops Research Institute for the Semi AridTropics, Patancheru, Andhra Pradesh,
India. pp 427–436
Onim JFM (1981b) Effects of two population improvement methods on grain yield of pigeon pea composite
populations in Kenya. Euphytica 30:271-275
Onwuka GI (2006) Soaking, boiling and antinutritional factors in pigeon peas (cajanus cajan) and cowpeas (Vigna
unguiculata). Journal of Food Processing and Preservation 30: 616-630.
Pal D, Mishra P, Sachan N, Ghosh AK (2011) Biological activities and medicinal properties of Cajanus cajan (L)
Millsp. Journal of Advanced Pharmaceutical Technology and Research 2:207-14.

Pande S, Sharma M, Naga Mangla U, Ghosh R, Sundaresan G (2011) Phytophthora blight of Pigeonpea [Cajanus
cajan (L.) Millsp.]: An updating review of biology, pathogenicity and disease management. Crop Protection 30:
951-957
Pande S, Sharma M, Guvvala G (2013) An updated review of biology, pathogenicity, epidemiology and management
of wilt disease of pigeonpea (Cajanus cajan (L.) Millsp.). Journal of Food Legumes 26:1-14.
Patel DB, Barve DM, Nagar N, Metha AR (1994) Regeneration of pigeonpea, Cajanus cajan, through somatic
embryogenesis. Indian Journal of Experimental Biology 35: 740–744.
Pathak GN (1970) Red gram. In: Pulse crops of India. Indian Council of Agricultural Research (ICAR), New
Delhi, India, pp 14–53.
Phatak SC, Nadimpalli RG, Tiwari SC, Bhardwaj HL (1993) Pigeonpeas: potential new crop for the southeastern
United States. In: J. Janick and J.E.Simon, editors, New Crops.Wiley, New York, pp.597–599.
Prasad S, Prakash R Haque MF (1977) Floral biology of pigeonpea. Tropical Grain Legume Bulletin No.7: 12.
Pundir RPS and Singh RB (1985) Cytogenetics of F1 hybrids between Cajanus and Atylosia species and its
phylogenetic implications. Theoretical and Applied Genetics 71:216–220
Pundir RPS and Singh RB (1986). Inheritance pattern for pod length and ovule number in wide crosses among
Cajanus and Atylosia species. Euphytica 35: 649–652.
Pundir RPS and Singh RB (1987) Possibility of genetic improvement of pigeonpea (Cajanus cajan (L.) ILLSP.)
utilizing wild gene sources. Euphytica 36: 33-37.
Ramanujam S, Singh SP (1981) Pigeonpea breeding in the allIndia coordinated program. In: International Workshop
on Pigeonpeas, vol 1. 15–19 December 1980, International Crops Research Institute for the Semi Arid Tropics,
Patancheru (India). pp 403–414.
Ramanadam P (1990) Pigeonpea: genetic resources. In: The pigeonpea (Nene YL, Hall SD, Shelia VK eds), pp
349–374. CAB International, Wallingford, Oxon, UK.
Rao KN, Reddy LJ, Bramel PJ (2003) Potential of wild species for genetic enhancement of some semi-arid food
crops. Genetic Resources and Crop Evolution 50:707-721.
Rao KS, Sreevathava R, Sharma PD, Keshamma E, Udaya KM (2008) In planta transformation of pigeon pea: a
method to overcome recalcitrancy of the crop to regeneration in vitro. Physiology and Molecular Biology of Plants
14:321–328.
Reddy LJ, Green JM, Sharma D (1981) Genetics of Cajanus cajan (L.) Millsp. × Atylosia species. In: Proceedings
of the International Workshop on Pigeonpeas, Volume 2, pp 39-50, ICRISAT, Patancheru, Andhra Pradesh, India.
Reddy LJ and Mishra AK (1981) Is emasculation necessary for pigeonpea hybridization? International Pigeonpea
Newsletter 1:12–13.

Reddy SJ and Virmani SM (1981) Pigeonpea and its climatic environment. In: Proceedings of the International
Workshop on Pigeonpeas. Vol .1, pp 259-270, 15-19 December 1980, ICRISAT, Patancheru (India).
Reddy LJ (1990) Pigeonpea: Morphology. In: The pigeonpea (Nene YL, Susan DH, Sheila VK Eds.), pp 47-87.
CAB International, UK.
Reddy LJ, Saxena KB, Jain KC, Singh U, Green JM, Sharma D, Faris DG, Rao AN, Kumar RV and Nene YL (1997)
Registration of high protein elite germplasm ICPL 87162. Crop Science 37: 94.
Reddy LJ, Kameswara Rao N, Saxena KB (2001) Production and characterization of hybrids between Cajanus cajan
× Cajanus reticulates var. grandifolius. Euphytica 121:93–98
Rekha R, Prasanthi L, Reddi Sekhar M, Latha P, Sudhakar S (2011) Genetic diversity in pigeonpea [Cajanus cajan
(L.) Millsp]. Legume Research 34: 139 – 142.
Remanandan P, Sastry DVSSR, Mengesha MH (1988) ICRISAT pigeonpea germplasm catalogue: evaluation and
analysis. Patancheru, Andhra Pradesh 502324, India: International Crops Research Institute for the Semi - Arid
Tropics. 95 pp.
Roy A and De DN (1965) Inter-generic hybridization of Cajanus and Atylosia. Science and Culture 31: 93.
Roy B (1933) Studies in the development of the female gametophyte in some leguminous crop plants of India.
Indian Journal of Agricultural Science 3: 1098.
Royes WV (1976) Pigeonpea. In: Evolution of Crop Plants (Sommonds NW Ed.). 154–156 (Longmans, London
and New York).
Sardana S, Singh M, Sharma SK, Rajan N (2011) Plant genetic resources and conservation of biodiversity. In:
Biology and breeding of food legumes (Pratap A, Kumar J eds.). CAB International, UK, pp. 362-375.
Saxena KB, Singh L, Gupta MD (1990) Variation for natural outcrossing in pigeonpea. Euphytica 46:143–148.
Saxena KB, Chauhan YS, Johansen C, Singh L (1992) Recent developments in hybrid pigeonpea research. In: New
Frontiers in Pulses Research and Development. In: Proceedings of National Symposium, 10-12 November 1989,
pp 58-69, Kanpur, India, Directorate of Pulses Research.
Saxena KB, Reddy LJ, Singh L, Kumar RV and Faris DG (1998) Registration of ICPL 87154, a partially cleistogamous
pigeonpea germplasm with low natural out-crossing. Crop Science 38: 556.
Saxena KB (2005) Pigeonpea. In: Genetic Resources, Chromosome Engineering, and Crop Improvement (Singh
RJ, Jauhar PP eds.), pp 86-115, CRC Press, Taylor and Francis, New York.
Saxena KB (2006) Seed production systems in pigeonpea. pp 76. International Crops research Institute for the Semi
Arid Tropics, Patancheru, Andhra Pradesh, India.
Saxena KB (2008) Genetic Improvement of Pigeonpea — a review. Tropical Plant Biology 1:159–178.

Saxena KB, Kumar RV (2010) Insect-aided natural outcrossing in four wild relatives of pigeonpea. Euphytica 173:
329–335.
Saxena KB, Kumar RV, Sultana R (2010a) Quality nutrition through pigeonpea–a review. Health 11:1335–1344
Saxena KB, Nadarajan N (2010) Prospects of pigeonpea hybrids in Indian agriculture. Electronic Journal of Plant
Breeding 1: 1107-1117.
Saxena KB, Sultana R, Mallikarjuna N, Saxena RK, Kumar RV, Sawargaonkar SL,Varshney RK (2010b) Male-
sterility systems in pigeonpea and their role in enhancing yield. Plant Breeding 129: 125–134.
Saxena RK, Prathima C, Saxena KB, Hoisington DA, Singh NK, Varshney RK (2010c) Novel SSR markers for
polymorphism detection in pigeonpea (Cajanus spp.). Plant Breeding 129:142–148
Saxena RK, von Wettberg E, Upadhyaya HD, Sanchez V, Songok S, Saxena KB, Kimurto P, Varshney RK (2014)
Genetic Diversity and Demographic History of Cajanus spp. Illustrated from Genome-Wide SNPs. PLoS One 9:
e88568.
Sharma D, Green JM (1980) Pigeonpea. In: Hybridization of crop plants (Fehr, Walter R, Hadley, Henry H eds.),
pp 471 – 481, American Society of Agronomy and Crop Science Society of America, Madison, USA.
Sharma PC, Gupta PK (1982) Karyotypes in some pulse crops. Nucleus 25: 181.
Sharma KK, Lavanya M, Anjaiah V (2006) Agro-bacterium-mediated production of transgenic pigeonpea (Cajanus
cajan L. Millsp.) expressing the synthetic BT CRY1AB gene. In Vitro Cellular & Developmental Biology - Plant
42:165–173.
Sharma M, Rathore, A, Mangala UN, Ghosh R, Sharma S, Upadhyaya HD, Pande S (2012) New sources of
resistance to Fusarium wilt and sterility mosaic disease in a mini-core collection of pigeonpea germplasm. European
Journal of Plant Pathology 133: 707-714.
Sharma M, Telangre R, Ghosh R, Pande S (2015) Multi-environment field testing to identify broad, stable resistance
to sterility mosaic disease of pigeonpea. Journal of General Plant Pathology 81:249-259.
Sheldrake AR (1984) Pigeonpea. In: The physiology of tropical field crops (Goldsworth PR, Fisher NM Eds.), pp
385-417, John Wiley and Co, Chichester, USA.
Sheldrake AR, Narayanan A (1979) Growth develop-ment and nutrient up take in pigeonpea (Cajanus cajan).
Journal of Agricultural Sciences (UK) 92: 513-526.
Singh AK, Rai VP, Chand R, Singh RP, Singh MN (2013) Genetic diversity studies and identification of SSR
markers associated with Fusarium wilt (Fusarium udum) resistance in cultivated pigeonpea (Cajanus cajan). Journal
of Genetics 92: 273–280
Singh F, Singh IP, Mazumder ND (2011) Identification of Fusarium wilt-resistant sources of long-duration pigeonpea
(Cajanus cajan). Indian Journal of Agricultural Sciences 81: 1046-1051.

Singh NK, Gupta DK, Jayaswal PK, Mahato AK, Dutta S, Singh S, Bhutani S, Dogra V, Singh BP, Kumawat
G, Pal JK, Pandit A, Singh A, Rawal H, Kumar A, Prashat RG, Khare A, Yadav R, Raje RS, Singh MN, Datta
S, Fakrudin B, Wanjari KB, Kansal R, Dash PK, Jain PK, Bhattacharya R, Gaikwad K, Mohapatra T, Srinivasan
R,Sharma TR (2011) The first draft of the pigeonpea genome sequence. Journal of Plant Biochemistry and
Biotechnology 21:98–112.
Singh U (1998) Anti-nutritional factors of chickpea and pigeonpea and their removal by processing. Plant Foods
for Human Nutrition 38: 251-261
Sinha SSN, Kumar P (1979) Mitotic analysis in thirteen varieties of Cajanus cajan (L.) Millsp. Cytologia 44: 571.
Sivaramakrishnan S, Seetha K, Reddy LJ (2002) Diversity in selected wild and cultivated species of pigeonpea using
RFLP of mtDNA. Euphytica 125:21–28.
Spence JA, Williams SJA (1972) Use of photoperiod response to change plant design. Crop Science 12:121–122.
Sundaraj DD, Thulasidas G (1980) Botany of field crops. Pp 496. The Macmillan Company of India Ltd, New Delhi.
Surekha C, Arundhati A, Seshagiri Rao G (2007) Differential response of Cajanus cajan varieties to transformation
with different strains of Agrobacterium. Journal of Biological Sciences 7:176–181
Thiruvengadam V, Muthiah A (2007) Inter-specific hybridization between Cajanus cajan and Cajanus cajanifolius.
Crop Breeding and Applied Biotechnology 7:204–11.
Thu TT, Mai TTX, Dewaele EFS, Tadesse Y, Angenon G, Jacobs M (2003) In vitro regeneration and transformation
of pigeonpea [Cajanus cajan (L.) Millsp]. Molecular Breeding 11:159–168.
Tuteja R, Saxena RK, Davila J, Shah T, Chen W, Xiao YL, Fan G, Saxena KB, Alverson AJ, Spillane C, Town C,
Varshney RK (2013) Cytoplasmic male sterility-associated chimeric open reading frames identified by mitochondrial
genome sequencing of four cajanus genotypes. DNA Research 20: 485–495.
Upadhyaya HD, Reddy KN, Gowda CLL, Singh S (2007) Phenotypic diversity in the pigeonpea core collection.
Genetic Resources and Crop Evolution 54: 1167-1184.
Upadhyay B, Parveen, Dhaker AK, Kumar A (2010) Ethnomedicinal and ethnopharmaco - statistical studies of
Eastern Rajasthan, India. Journal of Ethnopharmacology 129:64–86.
Upadhyaya HD, Reddy KN, Pundir RPS, Singh S, Gowda CLL, Ahmed MI (2013) Diversity and geographical
gaps in Cajanus scarabaeoides (L.) Thou. Germplasm conserved at the ICRISAT genebank. Plant Genetic Resources
11 (1), 3–14.
Van der Maesen LJG (1980). India is the native home of the pigeonpea. In: Libergratulatorius in Honerem HCD
de Wit (Arends JC, Boelama G, de Grant CT, Leeuwenberg AJM Eds.), pp 257–262, Agricultural University
Miscellaneous Paper, vol 19, Wageningen, The Netherlands.
Van der Maesen LJG (1986) Cajanus DC. and Atylosia W. & A. (Leguminosae), Agricultural University Miscellaneous
Papers 85–4 (1985). Agricultural University, Wageningen, The Netherlands.

Van Den Beldt RJ (1988) Cajanus cajan: it is more than just a pulse crop. NFT Highlights, NFTA, 88-06: 1-2.
Van der Maesen LJG (1990) Pigeonpea: origin, history, evolution and taxonomy. In: The Pigeonpea (Nene YL,
Hall SD, Sheila VK eds.), pp 15-46, CAB International, Wallingford, UK.
Varshney RK, Penmetsa RV, Dutta S, Kulwal PL, Saxena RK, Datta S, Sharma TR, Rosen B, Carrasquilla-Garcia
N, Farmer AD, Dubey A, Saxena KB, Gao J, Fakrudin B, Singh, MN, Singh BP, Wanjari KB, Yuan M, Srivastava
RK, Kilian A, Upadhyaya HD, Mallikarjuna N, Town CD, Bruening GE, He G, May GD, McCombie R, Jackson
SA, Singh NK, Cook DR (2010) Pigeonpea genomics initiative (PGI): an international effort to improve crop
productivity of pigeonpea (Cajanus cajan L.). Molecular Breeding, 26: 393–408.
Varshney RK, Chen W, Li Y, Bharti AK, Saxena RK, Schlueter JA, Donoghue MTA, Azam S, Fan G, Whaley AM,
Farmer AD, Sheridan J, Iwata A, Tuteja R, Penmetsa RV, Wu W, Upadhyaya HD, Yang SP, Shah T, Saxena KB,
Michael T, McCombie WR, Yang B, Zhang G, Yang H, Wang J, Spillane C, Cook DR, May GD, Xu X, Jackson
SA (2011) Draft genome sequence of pigeonpea (Cajanus cajan), an orphan legume crop of resource-poor farmers.
Nature Biotechnology, 30:83–89.
Varshney RK, Kudapa H, Roorkiwal M, Thudi M, Pandey MK, Saxena RK (2012) Advances in genetics and
molecular breeding of three legume crops of semi-arid tropics using next-generation sequencing and high-throughput
genotyping technologies. Journal of Biosciences 37:1–10.
Varshney RK, Murali Mohan S, Gaur PM, Gangarao NVPR, Pandey MK, Bohra A, Sawargaonkar SL, Kimurto PK,
Janila P, Saxena KB, Fikre A, Sharma M, Pratap A, Tripathi S, Datta S, Chaturvedi SK, Mallikarjuna N, Anuradha
G, Babbar A, Choudhary AK, Mhase M B, Bharadwaj Ch., Mannur DM, Harer PN, Guo B, Liang X, Nadarajan
N, Gowda CLL (2013) Achievements and prospects of genomics-assisted breeding in three legume crops of the
semi-arid tropics. Biotechnology advances 31: 1120–1134.
Vavilov NI (1951). The origin, variation, immunity and breeding of cultivated plants. Chronica Botanica 13: 1- 366.
Williams IH (1977) Behaviour of insects foraging on pigeonpea (Cajanus cajan (L.) Millsp.). Tropical Agriculture
54:353–363.
Yadav PBS, Padmaja V (2002) Interspecific hybridization and evolutionary relationships of (L.) Millspaugh and
four of the wild species. Cytologia 67: 67–73.

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1.1 Classification and Nomenclature ..................................................................... 1

1.2 Botanical Description of Pigeonpea ................................................................. 2

1.3 Economic Importance and Nutritional Composition....................................... 4

2. AREA, PRODUCTION AND PRODUCTIVITY........................................................ 5

2.1 Geographic Distribution.................................................................................. 5

2.2 Distribution in India including Regions of Cultivation ................................... 6

2.3 Zonalization and Varietal Testing System for Release........................................ 6

3. GEOGRAPHIC ORIGIN, GENOMIC EVOLUTION AND.................................... 7

3.1 Centres of Origin and Diversity....................................................................... 7

3.2 Genomic evolution.......................................................................................... 9

3.3 Genetic Diversity of Indian Germplasm........................................................... 10

3.4 Germplasm Collection at National and International Institutes........................ 11

4.2 Methods of Pollination, Known Pollinators and Pollen Viability...................... 11

4.3 Seed Production and Dispersal......................................................................... 12

4.4 Mating Systems................................................................................................ 12

4.5 Methods of Reproductive Isolation................................................................... 13

4.6 Potential of Vegetative Propagation.................................................................. 13

5.1 Intraspecific Crosses ........................................................................................ 13

5.2 Naturally Occurring Interspecific Crosses......................................................... 13

5.3 Experimental Interspecific Crosses.................................................................... 14

5.4 Genetic Introgression....................................................................................... 15

6. Human health implications.......................................................................... 16

7. Known interactions with other organisms in managed.............. 16

7.1 Interactions in Unmanaged and Managed Ecosystems...................................... 16

8.2 Sowing Season.................................................................................................. 20

8.3 Seed Production............................................................................................... 20

9.1 Milestones in Breeding..................................................................................... 22

9.2 Heterosis Breeding in Pigeonpea ..................................................................... 22

9.3 Major Traits of Interest and Priorities in Breeding............................................ 22

9.4 Advancements and Challenges ......................................................................... 23

10. Biotechnological interventions in pigeonpea................................. 24

10.1 Transgenic Approaches for Genetic Improvement............................................. 24

10.2 Genomics and Molecular Breeding in Pigeonpea ............................................. 25

1. INTRODUCTION The botanical name, C. cajan (L.) Millsp. has

substance on their surface. It is the second most Subtribe Cajaninae

Biology of Cajanus cajan L. (PIGEONPEA) 1

1.2 Botanical Description of Stem

Biology of Cajanus cajan L. (PIGEONPEA) 2

Biology of Cajanus cajan L. (PIGEONPEA) 3

Biology of Cajanus cajan L. (PIGEONPEA) 4

2. Area, production and productivity

Biology of Cajanus cajan L. (PIGEONPEA) 5

Pigeonpea is a photo thermo sensitive crop and

Biology of Cajanus cajan L. (PIGEONPEA) 6

For the purpose of varietal release, elite pigeonpea

1. North Hill Zone (NHZ) comprising

3. Geographic origin, genomic evolution and

Biology of Cajanus cajan L. (PIGEONPEA) 7

Biology of Cajanus cajan L. (PIGEONPEA) 8

Biology of Cajanus cajan L. (PIGEONPEA) 9

Biology of Cajanus cajan L. (PIGEONPEA) 10