PLANT SCIENCE MONOGRAPHS

Edited by
Professor Nicholas Polunin

SEED PRESERVATION AND LONGEVITY

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PLANT SCIENCE MONOGRAPHS

Uniform with this volume will be:

4Biology ofMycorrhiza .
Encyclopdia of Weds and their Cfmtrol
Grassland Impro1Jtment
!!faT/groves o/the World.
Microbiology of the Atmosphere
Mutations and Crop Improvtml!l1t
4Plant Growth Suhstallf;es
Plallt Lift alld Nitrogen .
Salt Marshes and Salt Deserts of the rVorJd
SeJ. in the Lower Organisms

J. L. Harley
L. J. King
A. T. Semple
V. J. Chapman
P. H_ Gregory
A. Gustafsson
L. J. Audus
G. Bond
V. J. Chapman
H. P. Papazi:m

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FURTI-fER TITLES ARE l)1\<"DER CONSIDERATION

THE feeding and clothing of the world's teeming millions can continue to keep
abreast of population increases through the help of effective application of
research in the plant sciences. The publication of this research, by which means
a scientist or tcchnologist makes his findings known to workers elsewhere,tends to be scattered in literally hundreds of botanical and agricultural journals
emanating from most of the countries of the world. Often it appears in such
polyglot arrays of fragments that it is extremely difficult to bring together even
in some narrow 'line' of endeavour. Consequently advances are slowed and interests unnecessarily divided, scientific and human progress being thereby
retarded.
The present scries of 'monographs' is designed to remedy these deficiencies
in especially important or attractive specialities, by publishing individual booklength accounts of the entire background and current progress in their fields.
Such detailed surveys, being fully documented and plentifully illustrated, should
prove of real value to the world at large in constituting the bases for further
advances on the ever-expanding'hoOzons of scientific research, and so lead to
improved productivity and, ultimately, standards of living. They are prepared
by spt"Cialists usually of international reputation for their work in the field
chosen, and often culminate a lifetime of active investigation. Being as up-todate as possible, they will often embody significant advances not previously
published.

Already published and :I.v";bble.

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.~, Relative r.lte of gromh of seedlings produced by freshly han-ested and old seeds of the
Sacred or Indian Lotus (NdlimbIJ nucif~,a).
D, Structure of seed-coat of Na"mbo sho,,~ng w\l{er_resist\lnt layen, n:nnely, (a) epidermis and
(b) pan of palisade layer to the 'light-line' (m), all greatly ma~ficd.
C. 30- to 6o-foot banl.: left by the CUt of the ri"er; seeds were located in 5tnlta 3 to 6 feet
from the top of the pre<.:ipice, as ;ndie-,lIed by arrow,;.
D, Plant grown from olle of the old seeds.

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PLAl\'T SCIENCE MONOGRAPHS

edited by
PIlOl-T..'lSOR NIOIQLMi PO!.UNIN

SEED PRESERVATION
AND LONGEVITY
By
LELA V. BARTON
M.A., Ph,D. (Co/umbill)
Boyce Thompsoll

rnstitut~

for Plant Rtuarch,

Yonkus, Nt11J Yo,k, U.S.A.

'96 ,
LONDON

3 5 527 -

LEONARD HILL [BOOKS] LIMITED

L\lTERSCIENCE PUBLISHERS, INC.
NEW YORK

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LONDON
LeoIl:lrd Hill [Books] LId.

9 Eden Strcct, N.W.!

NEW YORK
Intc.rscicnce Publishers Inc.
250 Fifth Avenue, New York I

FIRST PUDUSHED IN GREAT BRITAIN 196r

copyright Lcb V. manon, J96r

l'!UNITD L'I GR.EAT 1lR1TlllN ;\T

ll'f lINIYl;RSln" I'Ilf.SS
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INTRODUcrION
Ol\c: of the most important factors contributing to ma.~imum agriculturnl
crop production per unit of land is tbe use of good seeds. Such seeds
should haye a high purity v-alue, i.e. they should be free from all other seed
types, whether of weeds or of cultivated plants, and their gcnninarion
capacity and vigour should be high, thus providing tht: best possible
guarantee of a good seedling stand-in spite of diseases, insects, com

petition of weeds, and unfavourable weamer conditions. This fact has, of

course, ~n recognized for a long time; but the systematic storage of
seeds to IruLinrain viability, and the routine testing and analysis of seeds
fOT

planting value, dates from only about the beginning of this century.

Jncreased interest in, and knowledge of, the seed problem have resulted
in the establishment of private, state, and federal seed-testing laboratories
and the formation of national and international rules for seed testing.
Along with improycd methods of testing has come the necessicy for proper
storage facilities to maintain yiabiliry'. h is importlUt [Q seedsmen to be
able to keep surplus supplies for sale in bter years. In some instances this
has been done without the benefit of proper storage conditions for
maintaining high-quality seeds. As a result, everyone who buys seeds is
apt to get lots which arc of low quality or which fail to gcrminate. As morc
information is secured on the storage requirements, reputable secdsmen
are building dehumidified or cold-storage rooms. The possibility ofholding
seed" in storage for at least two years reduces the amount ofland needed
for seed production, as crops can then be alternated.
Many coniferous trees do not set seed eyer')' rear: indeed there mar be
an interval of as much as ten rears between crops. This poses a serious
problem in rcafforestation projects which dcpend on a constant supply of
seeds for nursery usc. Also, direct reseeding of forest land could be
seriously hampercd by the lact.: of a seed supply in any given ycar. With
the benefit of below-frcezing stonfge, it is safe to hold conifer seeds for
periods adequate [Q cover these needs. Such low temperatures automaticUly control insect damage, which is considerable in many seeds. Insect
control has usually depended on the presence of certain insecticides which,
howevCf, introduce the funber problem of injury [Q the seeds or to the
pasons handling the seeds.
The possibility of long-term storage is a boon to persons concerned
.. It h:Ls not been found necessary to distinguish in this work
sud-like fruits (Ed.).

betWCCIl

seeds and

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SF.F,D PRFSf>RVATION A,-";D LONGEVI'I'Y

with generics or with plant introduction, for it provides a constant source

of valuable seed stock \yithout the necessity of growing large numbers oL
plants and controlling seed production-a laborious and difficult process.
Granted that good storage conditions for a cerrain seed have been
determined, there are still other questions which arise. '~lin the seeds from
storage, especially low-temperature storage, survive removal for packeting,
which must be done well in advance of tne rime the consumer buys and
plants them? Furthermore, after packeting, seeds are sometimes stored
under very unfavourable conditions on the retailers' shelves. Some
experimental results on this problem have demonstrated the efficacy of
moisture-proof packets. The best storage condition, determined for a
specific seed, prandes it with maximum resistance to harmful conditions
following packeting for sale. Additional work needs to be done .in this
ficld, using different kinds of seeds; but the principles already determined
will-probably' apply to many species.
The quality of plants which can be expected from old seeds is another
matter of importance. An increasing body of evidence shows that the
performance of any seed depends less upon its age than upon the conditions under which it has been stored.
The absolute dependence of the world's population on plants for food,
and the realization that most plants are grown from seeds, should be
sufficient to dramatize the importance of conserving our seed supply.
A basic bibliography of literature on the subject is to be found in seyeral
general articIe.<; or books (Franck, 1928; Crocker, 1938; Barton & Crocker,
1948; Crocker, 1948; Crocker & Barton, 1953; and Owen, -.1956).

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PREFACE
Spama{OpJryles, or seed plants, arc morc important to man than any other
liying plants, for they are best adapted to cultivation and furnish, directly
or indirectly, most of the food and clothing for the entire world. These
phnts are distinguished from lower groups by the production of seeds. Dy
the development and establishment of the seed habit, the reproduction of

the individual plant and the continuance of the species arc assured.
Furthermore, the seed is an excellent method for multiplication and
distribution of the species. The seed plants are divided into Gyml/osperms
;md A1lgiosperms. The former produce 'naked' seeds, which arc so called
bec1usc they are borne on the surface of a scale, usually of a cone, as in

pine (Sl"e' Figure t, A'). Gymnosperms are woody perennial forms comprising a group of about SOO species, most of which are evergreens.
They include many commercially important types that arc valuable for
timber and resins of various kinds.
In the Angiosperms, the seeds are borne in an enclosed structure or
L"'1!it developed from the ovary. This fruit may be dry or fleshy, and may
mnuin one or several seeds. The Angiosperms dominate the modern plant
_-arId, being represented by some 250,000 species. They are divided into
[lIfO groups, MOllocolyledollS and Dicotyledo1lS. There are several structural
dilferences between these two groups; but, from the point of view of seed
distinctions, the former possess embryos with only one terminal cotyledon,
bile the embryos of the latter have two lateral cotyledons. All grasses,
iDcluding the commercially important cereal and many forage crops, are
monocotyledonous, while mOst of the vegetables, flowers, and deciduous
belong [Q the Dicotyledons.
Seeds are usually produced as a result of pollination and fertili7.ation.
~g is the first step in such seed production. In the ordinary way,
.am., produced in great abundance in the anthers of the stamens-the
m.Jc parts of the flower-is carried by wind, insects or other means to the
~ which is the tip of the pistil or female part of the flower. Here the
.-en is commonly held by a special secretion in which the pollen grain
_'O.mitutes to produce a rube which carries the male nucleus down the
st\k C!f me pistil to the ovary, which contains the ovules. The pollen tube
f"''''~ues the ovule and normally its germ nucleus fuses with the egg
~ of the ovule to form the embryo of the seed.
.Embryos vary greatly in size and appearance, but, v;ith few exceptions,
drJ .L~ m3de up of the same organs: a plumule or rudimentary shoot,
.. T......;.,nV,llcrms, ctc. arc explained in the Glossary commencing on p. 162.
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SEED PRF.5ERVATION .o\...1\'D LONGE\'ITY

cotyledons or seed-leaves, hypocotyl between the cotyledons and ~lumulc-,

and the radicle or rudimentary root. The plumulel hypocotyl,
rad.icle
fann the axis of the embryo. Jn Dicotyledons the cotyledons may rCrrulin
underground upon germination of the set:d, or they may be pushed above
ground by the growth of the hypocotyl to form fleshy green le:lYes as in
bean. The single cotyledon (scutellum) of grasses (lHonocotyledons)
remains as a food-absorbing organ wilhin the germinating grain. Iruome
seeds, as for example those oforchid'ior holly, the embryo is an undifferentiatcd mass of cells at the time the seed matures in the plant. Such
embryos must develop the structures described above before the~" arc
ready to form a new plant. This results in delayed germin..1.tion when the
seed is planted.
The endosperm, or food stored fOr use by the young embryo, also
commonly develops in the seed following the conjugation of:l second male
nucleus from the pollen tube with two other nuclei in the ovule. This
means a double fertilization, one resulting in the young plant or embryo
and the other in the development offood for the young plant. Such double
fertilization occurs in most Angiosperms, and results in the form:ltioRof
the endosperm, a definite structure containing stored food. However,
the endosperm is not always present in the mature seed, as it mar have
been used up during maturation of the seed. In the bean seed, for example,
most of the stored food has been absorbed by the cotyledons which grow
to fill most of the seed cavity. Maize, wheat and other cereals, on the other
hand, have abundant endosperm.
JUSt as the sced is the ripened ovule composed of embryo and seedcoats together "~th stored food, so the fruit in which the seed is borne is
the ripened ovary together "itb other parts of the flower whieh may have
become associated with it. There are many different kinds of fruits, but,
for the present purpose, it is perhaps sufficient to classify them as simple,
aggregate, accessory, or multiple.
Simple fruits, or those resulting from the ripening of a single ovary,
may be further classified into dry or fleshy. Dry fruits are dehisccut or
indehisc6nt. Ivlany-seeded dehiscent fruits include pods and capsules of
various kinds which open when dry to liberate the seeds within them. Pods
m:!.y upen through one suture (larkspur, columbine, miU.."Weerl); one or
two sutures (legumes, see Figure I D); a circular line cutting off the
upper part of the fruit as a lid (plantain); or separation of two valves of a
capsule (mustards). Indehi'icent-dry fruits arc mostly one-secded or have
very few seeds. These fnuts arc commonly known as 'seeds' because-they
do not open to discharge the true seeds, but adhere tightly to them. Oncof
the most common of these dry, indehiscent fruits is the grain, or caryopsis.
Here the single seed completely fills the fruit, as in ma17..c (see Figure r E)
and wheat, for example. In addition to the fmit-coats, grass (seeds' oftcn
have small bracts (the lemma and palea) of the original floret associatcd
with them. Other indehisccnt dry fruits are samaras or keys, which are

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PREFACE

provided with a wing for dispersal. Examples of this type arc the fruits of
ash, with a terminal wing, or of elm, where the wing forms a margin around
the entire fruit, or of maple (st't Figure, I c), which is a double samara
or pair of fruits conspicuously winged from the apex. Achenes are small
fruits, which often have appendages (dandelion, lettuce) to act as agents of
dispersal.. The actual fruits of the strawberry are also achencs-in this
case produced on a fleshy flower-receptacle which really has no connection
with the pistil or female part of the flowcr. A nut is another hard, onecelled and one-seedcd, indehiscent fruit (beech, chcsmut, oak).
Fleshy fruits which are also indemscent may be fleshy throughout,
with or \\-ithout a firm rind or shell, Or lhc)' may be fleshy e.xtcmally and
hard or stony internally. Among the latter type are drupes or stone fruits
such as those of cherry or peach. The pomes (apple, pear, quince) are
fleshy fruits composed of two or several carpels of bony or cartilaginous
texture enclosed in flesh which morphologically belongs to the calyx and
receptacle of the flower from which the fruit was produced (sec Figure
1, R, B1). The fruit of the gourd and squash is a pepo, a fleshy fruit with a
hard rind. Other examples of simple fleshy fruits arc the hesperidium, a
special type of berry with a leathery rind represented by fruits of orange,
lemon and lime, and the berry itself which includes the fruits of grapc,
currant, banana and tomato, in which the fruit is l1e.";hy throughout.
Aggregate fruit.... are those in which a cluster of pistils, all from one
flower, is crowded on the Oower receptacle in one mass, as in raspberry
and blackberry. Multiple or collecti\+c fruits, on the other hand, arc those
which result from the aggregation of several flowers into one mass (pineapple, mulberry, fig).
The seed industry, one of the most important in the world because it is
at the heart of the survival of man, involves Yaried investments and skills.
It requires the senices of geneticists and plant breeders to develop and
select Yarietics superior in performance and in yield of vital food and
clothing materials. Plant physiologists are constantly working to determine
the effect of nutrition and environmental conditions and many other
factors on growth, seed-set and quality. Plant pathologists aim to control
the diseases of plants, and entomologists study the insects which affect
plant life in a never-cnding battle to reduce the ravages of these pests.
Growers must practise the highly specialized techniques of seed production. Special attention must be paid to the harvest and curing of seeds.
Germination problems arc many, and they must he determined for each
seed type. Testing and analyses of seed-lots have become problems of
major concern to every country, and most countries now have set certain
<otanclards which must be met before seed can be offered for sale in them.
Seed marketing forms an industry in itself. Finally, equipment manufxturcrs must provide the apparatus and materials required for the
functioning of this comple.'t industry.
Some idea oftheenent ofme seed industry in the world may be obtained
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SUI) PRfSI::In'AT10N Al'ID LONGEVITY

from a few statistics of seed production in the United States of America.

In H)S6, 75,950,000 acres of maize (U.S. corn) were han'csted to yield
3,45 1 ,292 ,000 bushels of grain; wheat h:J.n-cstcd from 49,817,000 acres
prouuced 997,207,000 bushels of seed. In 1955, ::212,390,000 lb. of clean
alfalfa seeds valued at $43,752,340 were grown on 1)92,500 acres.
During the same year 204,757 acres were planted in vegetables which
yielded 216,294,000 lb. of seeds. The bean crop alone in 1955 required
1,5 02 ,000 acres to produce 1,664,900,000 lb. with a market value of
51 16,79,490 .
In addition to the seeds grown for the production of food, clothing,
and shelter, for man and domesticated animals, there is a large and
expanding flower-seed industry. Tree seeds also add their importance to
those already described. Tons of such seeds arc needed each 1"car for
rcalforestation of lands throughout the world.
Members of the seed industry have the grave responsibility of furnishing a good start for gro\\"th of important crops by proy:iding seeds of
desirable strains and high quality, year after year. It is always necessary
to hold seeds from the time of harvest until planting time, and :it :is often
desirable, in the interest of economy, to store excess ban'cst for one. or
several seasollS. It becomes neeessa~y then to know safe storage conditions, which vary from seed type to seed type. Also, safe shipment of
fresh seeds :is often a problem---especially if the)' musr be- routed through
hot, humid climates. Over the years, seed p~'siologists and other individuals concerned havc- tried to uncover some of the secrets held wirhin
the coats of seeds, both viable anti dead, in order to determ:ine their
keeping qualities under various conditions.
This book is an attempt to combine in one volume the available
information concerning the longevity ofseeds under natural and controlled _
conditions. The scientific aspects of the problem are considered in so
far as they arc known, and the practical applications of the existing
knowledge are pointcd out. No attempt has been made to list all references
to published articles on the general subject of 'Seed Preservation and
Longevity', for this would be tedious and repetitiolls. R"1ther, literature
has bcen selected which, in the opinion of the author, is TeprCSCntative
of the various phases: that should be covered in such a work.
The author is grateful to Dr. George L.A'leNew, Managing Director
of the Boyce Thompson Institute for Plant Research, Inc., for his
interest and help; to Mrs. Jean G. Fine, Miss Joyce Fitzgeorge, and Dr.
Norma E. Pfeiffer, for the checking of literature and for proof reading; and
to Messrs. L. P. Flory and "iilliam G. Smith, Jr., for photograph:ing and
arranging platcs. The author is also grateful to the follo"'ing for perm:i~ion
to reproduce certain figures from original research articles: Drs. G. C.
Papavizas and C. M. Christensen, Fig. 7 and Pute 2; Drs. N. Armolik,
J. G. Dickson, and A. D. Dickson, Plate 3; Associared Seed Growers,
Inc. Phtes 6 and 7; Dr. G. Lakon, Figs. 12 anu13; Dr.!,;,1. Simak, Plate 9;

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I'RU'ACE

Drs. E. 11. Toole, V. K. Toole, and H. A. Borthwick, Pbte 10, b; and

Drs. II. Gunrhardt, L. Smith, M. E, Haferkamp, and R. A. Nilan, Plate '3
An acknowledgment is given in the legend for each of these figures.
Acknowledgments for permission to reproduce tables from various
publications are also gi\'cn in the text.

l..EL.o\ V. B.o\RTON
llon:l' TlfO~lf'SOS' L"'IST1Tt.JT! FOR Pl.M\'T RESf_'LROl, L"c..

Yo.''KERS, NEW YQRX:

CONTENTS
v

INTRODUCTION

vu

PREFACE

LIST OF TABLES
LIST OF Tu..USTR.HIONS
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OLD SEEDs.

Egyptian Mummy Wheat

I RECORDS OF

II LIFE-SPAN OF SEEDS BURIED IN SOiL

Moist Storage of Seeds of Amarflllf!lIIs relrofll!xlIs L

ill MOISTURE En'ECTS

rI

14

Moisture Relationships of Seeds Stored 'Open' in the Laboratory

Artificial Drying

TY OTHER FAcroRS AFFECTING

Temperature Effects

22

28

30
30

LOl>:QEVITY

Gas Effects .

33

Light Effects

36

Inheritance
Sccd-eoat Effects
Maturity Effects
Dormancy Effects
Fruit or Pulp Effects

36
37
37
38
39

V 0rHER FACTORS AFFECTING

Microflora
Olemical Effects

(coutd.)

40
40
46

\'1 LONGEVITY OF VEffi.'"ABLE Al\'D FLO<;\'ER SEEDS.

SO

LONGEVITY

Vegetable Seeds
Flower Seeds

Vll

LONGEVITY OF SEEDS OF

50
S4

FruD

CROPS

63

Wheat.
~1aize .
Other Cereals

63
65
67

Gras.<;e.<;
Legumes

67
69

~1I1 LONGEVITY OF 01HER ECONO:\llC..u..LY IMPORTAh'T SEWS

Oily Seeds
Other Seeds .

7'
7'
75

Xlll

SEfD PRESERVATION A.I\,""D LONGEVITY

auPro<

IX

LONGEVITY OF

""
i8

TRf:f. StlTIS

Conifers

83
86
8i
8i
8i
88
88

SUDS elf-' AQmTIc Pt.Al\;l'S, ETC.

Seeds Injured by Desiceuion .

9'

8<)

LOXGEVITY OF

XI EFFECT OF P.\CKf"I'S oX' SED>

XlI

i8

Elms .
Ashes .
Willows
Birches
Poplars, etc.
Soulhern-Bccches .
Fruit Trees
Q!linine

91

VIARU.!TY

99

METHODS OF TESTING FOR VI.'LlJlUTY

Q!licl: Viability Tests

Excised Embryo Technique
Plasmolysis .
Enzyme Activity .
Chemical Treatments
Fluorescence Measurements

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115,
115
IIi
120

Xln MI:.TIIODS OF TISTING FOR VURlUIT (comd.)

Teuazolium Salts .
Electrical Responses

X-R.ay Analysis

XIV J>uI..'TS ~"RO~I OLD SEIDS

Various Other Plants

xv

CAUSES OF DETERIORATION

Chemical Composition .
Mutations
XVI PRACTICAL CoNSIDERATIONS

Transportation
Storage Facilities
GLOSSARY
BIBLIOGRAPHY
Cot\,\'l::ltSION F.o\CfORS
SUBJECT INDEX
AunlOR INOF,X

210

LI ST OF TABLES
T.UIU.li

"A(if:

I Record of old seeds

II ResuilS of all t~ts to date in Re:ll's buried seed experiment

111 Amaranl1Jus rttrofltrus. '942 crop. Germination after \1lrious

periods .

IV

~toisture

12

contents of seeds siored at

VariOliS

tempentures and

relat1\"c humidities
V Germination percentages obuinctl from seeds stored at ,,-allow

temperatures and rclath"c humidities

15

19

VI Estimated maximum safe seed moisture contents for storage

VII Germination and moisture content of conifer seeds after storage at
sub-freezing temperatures.

31

V1 TT Germination of vegetable seeds on moist filrer-paper at controlled

temperatures after storage under various conditions

52

IX Regal Lily. Seedling production in soil in the greenhouse after

storage .
X SCt:dling production in the greenhouse from pine seeds stored

air-dry under various conditions

21

60
8I

X I Seedling production in the greenhouse from conifer seeds stored in

sealed conuinas at 3pprorimatdr ~.f0 C .
81
XII Seedling production in the greenhouse from ..' merinn elm seeds
stored under various conditions "
86
Xill Germination of stored onion seeds after p3cketing and further
storage
102-3
xn' I.ist of speci(S, "'ith litennrre references, which ha'"e been tested
113-14
by the rapid viability method
>"'V Resu.lts obtained "'ith rapid 'iabiliry test and staining with 2, 3, 5rriphcnrltcrrazolium chloride in determining the 'iability of
various seeds "
128-9
XVI Results obtained with germination tests, rapid viability test, ami
staining with 2, 3, 5-triphenyltcrrazolium chloride, in determining the '>iabiliry of variou.s pinaceous seeds .
" 130-1
XVII Results obtained with rapid \"i3bilitr test and staining with 2, 3, 5triphenyltetrazo'u-n chloride in determining the viability of
various rosaceous seeds
132
xvrn Comparison of results betw(.-en rapid viability and tetrazolium
tests
133
XLX Cumulative weekly totals of Aster flower-heads cut during the
gro....'ing-scason of t945
1+3
XX TOlal pepper fruit )ields in ounces for seed-lots and blocks for [he
144
entire season
XXI Total tomato fruit yields, in ounces, for seed-lots and blocks for
145
the entire sca."On
XXI I Total weights oflcttuce plants, in ounces., for seed-Iors and blocks 147
xv

LIST OF ILLUSTRATiONS
PLATES
Germinative ...igour of old Indian Lotus seeds .

FrOl/lispircl'

facing pat!
Germ damage (.'aused by inoculation of wheat scetl

48

Effect on barley seed germination of filtrate,~ of fungi

49

4- (0) Delphinium plants produced from old seeds

(b) Regal lily plants prodl1ced from seeds stored for four
five years

[0

j.

Pinys lada plants produced from five-yt:ar-old seeds

6.

Apparatus which may be used for making a seed moisturecontent rcst .

,,6

Germinating cucumber seeds stored for two to five years

in tin cans .

S.
9.
to.

DC\-dopmcnt of excised embryos on moist filter-~pcr

Radiographs of impregnation patterns of seeds of Pima
sylvestr;s
(0) Young seedlings of beans from famul1Ible and unfavourable

Tq-IS
:
126

storage

(b) Mature plants of beans from fa,'ourable and unfavourable

storage

1'27

IT.

Peppers harvested from plants grown from fresh and old seeds

140

12.

Growth of tomato plants from seeds of different ages

147

13.

Chromosomal aberrations oa:urring lD plants from aged

wheat seeds

Sf!

FIGURES IN TEXT
P!lGl';

fIGURE

1.

2.

3.
4.
5.

Some representative seeds and fruits

Graphs indicating moisture content of seeds after storage
Graphs indicating ~rmination after swrage
Graph showing moisture contents at various times of the
rear of seeds stored 'open' in the laborator1:
Uraphs shov.ing the effect of temperarure and method of
scaling on the yiabilit~; of stored dandelion seeds
:\"Vl

:-.:viii

'7
20

23

n_
6.

LIST 01' ILLUSTRATIONS

,,~

Graph indicating percentage germination of seeds ofDouglas

.fir and Sitka spruce after storage at sub-freezing temj)eratures

Graphs showing germination and germ damage of grains of

Willet wheat non-inoculated and inoculated with Aspergillus
calldidus, A. repe1lS. and A. restric/us

8.

Graphs indicating viability of annual delphinium seeds of

the 1924. 1925, and 1926 crops

Graphs indicating viability of perennial delphinium seeds of

the 1925 and 1926 crops

<0.

Graph indicating effect of storage of American elm seeds

upon germination.
Graphs indicating percentage germination of onion seeds
held for twenty-four months at 2S"C. and then stored 10

...
12.

packets or tin-cans for a further period

Excised embryos of cereals

'3

Median longitudinal section through a kernel ofmaize.

xvii

3'

4'
55
57
85

105

"3

,.

A'

-E
FIG. t.-Some reprcsmr:u:ivc seeds and fruits. A, A' Cone and winged secl of Sugar
Pinc (Pinus lamb~tillfl!J). B, a'. Pome and seeW of Pf:l~ (PjruJ (Ommu.il cultivu William
Bartlett). C, Leaf and double samar.!. contl.i.ni.ng seeds of Sugar .Maplc (ActT'SIM'buum).
0, Pods :md seeds of Garden Pe:t (Pisum stltiwm). 1';, Gnins of~ (Z(,:l map), the one
on the lcfl germinaring.
XYlU

RECORDS OF OLD SEEDS

Tl-IE life-span of seeds has been a matter of interest, conjecture, and
fantastic report, for more than a hundred years. Turner (1933), in his
review of literature on the subject, says that Theophrastus mentions that
'of seeds some have more vitality than others as to keeping' and 'in
Cappadocia at a place called Petra, they say that seed remains even for
forty years fertile and fit for sowing'. All of the early reports, including
those made in the first part of the present century, were concerned with
seeds of more or less established ages, which were found, for example, on
herbariwn spccimcus or in old cupboards. Vcry little ifanything was known
about the origin of these seeds and the specific storage conditions which
may have affected their longevity. Furthermore, methods used for
germination tests rna)' not have been, and probably were not, those which
were best for measuring viability. In spite of these handicaps, some
"aluable data were obtained.
Ewart (1908), in his treatise 011 the Longevity of Seeds, pointed out
that probably few sections of human knowledge contained a greater
number of contradictory, incorrect, and misleading observations than
prevail in the works dealing with seed Viability, and that knowledge of
the subject was still in an incomplete and indeed fTagmentary condition
at that time. On the basis of reports of others and some work of his own,
Ewart divided seeds into three biological classes according, as he said, to
their duration oflife under favourable conditions. These were short-lived
or <microbiotic' seeds, with a life-span not exceeding three years, 'mesobiotic' seeds which may live from three to fifteen years, and 'macrobiotic'
seeds which may retain viability for fifteen to over a hundred years. Under
the macrobiotic classification he listed 137 species in 47 genera of the
f.unily Leguminosae; 15 species in 8 genera of Malvaceae; 14 species in 4genera of Myrtaceae; 3 species in 2 genera ofNymphaeaceae; 3 species in
3 genera of Labiatae; 2 species in 2 genera of lrideae; and I species in
each of the families Euphorbiaccae, Geraniaceae, Goodeniaceae, Polygonaceae. Sterculiaccae, and Tiliaceae.
One of the earliest records of germination of old seeds is that of
Candolle (1846). Seeds of 368 species representing various families were
mUected. principally from the botanical garden of Florence in 1831, and
lIft'Te tested in 1846 when nearly fifteen years old. They had been kept in a
cabinet protected from high humidity and extremes of temperature. Only
A
,

SEED PRESERVATION AND LONGEHTI'

seventeen of these species germinated when the seeds were planted..in suil,
and only one of these, Do/iehus ul1guiculatus, 'li' showed more than 50 per
cent germination. Leguminosae (5 out of 10 species tried) and Malvaceae
(9 out of 45 species tried) accounted for q. of the- 17 viable lots of seeds.
The seeds tested represented 180 annuals, 28 biennials, 105 perennials, and
44 woody plants, together with 1 I unidentified seed-lots. The author
concluded that the figures obtained seemed to prove that woody species
preserve their viability longer than others, and that biennials, none of
which 'vere found to be viable, deteriorated most rapidly.

TABLE I
RECORD OF OUJ SEEDS

(from Becquerel, 1934, p. T66z)

Macrobiotic species

Date

S",,"

001Ited

growing
in 1906

Mmosa gkm~rala FarsI;. 1853 5 out of ro

M(/ilotus !l1!~4 Gueld.
1851 3
"
Astragalus 1114ssilimsis

,.

urn.

Cyti:,us auslria~us L.
Lal'atrra pSntd(H)!hia
Do;[
Diotka palldjl()ra Rusby
En:um lens L.
Trifolium anJens~ L.
uw:amn l(ucoupllala L.
Srcchys r.epetifalia-Desr.
Cytisus bijlo,.~ L'Herit.
Cassia bi<apsularis L.
Cassia multijuga Rich.

,'... ,
0

1843

,842
18{1
184
J838
1835
1829
1822

18J9
1776

,
,
,
,
,
,
3

"

"
"
"
"
"
"
"
"
"

Swl<
!p'owing
III 1934
50utofJO

8,

"
"
" ,
"
W
" 3
"w
0

Ddcnnined Probable
longcvity
longevity
(yr.)
(yr.)

" ,f

"

"

55

"
"

,."
,.

86

"

"
"

"
"w

" w

"

"

"

"
"

w
w
w

'"
"0

63

6,
93

65
68

99

155

77

"

u5
158

'99

Becquerel (1907) tested seeds of 500 species from the seed collection of
the Museum of Narural History.in Paris. The authenticated ages of these
seeds were known to be betw'een 25 and 136 years. Germinations were
obtained ITom four families; Leguminosae, Nymphaeaceae, Malvaccae,
and Labiatae. Twentyoftheseseedswerefromtwenty-eighttoeighty_scvcn
years old. Later, Becquerel (1934) made another test on the viability of
the old collections of secds which he had used in H)o6, with the results
shown in Table r. All of these seeds which germinated were of Leguminosae, except for Laviltera. (Malvaceae) and Stl1chys (Labiatae). The seeds
of Cassia 71lUIlijuga germinated after 158 years of storage-apparently

me

fie Authorities for scientific plant namcs, which may be needed to indiClte
precise sense in which the hncr arc employed, ~rc usually giVCll onlr when original
obscnaLions are t:Illlcerned, or where specified by thc authors of ciled works.-Ed.

...::;:

.::=

:ii

a==

::

RECORDS OF' OlD SEEDS

a rccord at that time. Turner (1933) found that seeds of A11lJryfJis

l.'f1fneraria and Trifolium striatum remained \iable for ninety years and
those of CytiSlls JCOpdriUS, A1e/ilotus alba, Lotus uligillosus, and Trifolium
pr(JII!tl~

for eighty-one years.

Schjelderup-Ebbe (J936) had access to two collections of old seeds in
the Botanical Museum of the University of Oslo. The larger and older
collection was that of N, G. Moe, Head Gardener of the Botanical
Gardcn from 1857 to 1892. It contained seeds collected mainly during
the years 1820 to 1892 as well as some older seeds. The second collection
was that of A. G. Dlytt, collccted mainly during the period of his professorship, 1880 to 1898. Sceds of the first collecllon were stored in strong paper
bags with the name of the species and date of collection written on the
outside. The second collection was stored in corked bonles with the species,
rime, and place of collection written on labels pasted on the bottles. The
germination tests were run during the rears J932 and 1933. so the ages of
the seeds tested ranged from approximatelr 34 to 112 years. The seeds
included Gymnosperms and Angiosperms; the latter group included
monocotyledons as well as dicotyledons. In all, 1,254 different batches of
seeds were tested and relatively few of these included duplication of
species. Our of these, seeds ofonly fifty-three species showed anr germinanon. The oldest living seeds found in these studies were those of Astragalus utriger, cighty-two rears old, which gave 6 per cent germination.
Thc author stated that he had added two families of plants to those
bearing macrobiotic seeds: Cannaceae, with Call1l0 pallicufata seeds
sixty-nine years old; and Thymelaeaeeae, with Daphlle l1uzereWIJ secds
thirty-five years old. Seed size appeared to be unimponant, as both vcry
small and very large seeds had lost their viability. The only report of an
older viable seed from stored collections was from those of the British
:\luseum (Anon., 1942), which recorded the germination of a seed of
."elumho after about 250 years of storage as well as of seeds of Albiuia
pdibrissin after 149 years.
Exell (1931) reponed that Robert Brown found 15Q-ycar-old viable
seeds of Nelumbo nuijera (syn. Nelumbium sjJ(ciosum), the Sacred or
It has, unfortunately, prm"ed impossible to determine the c.l:act age of these
Sdll1flbo'seeds' in the British Museum, where Mr. A. W. Excll now points out that
X. wUa may ha\e been the species imoh'ed. The seeds were from the Sloane colleelion, and concerning them J. E. Dandy (The Sloane Habarium, British Museum,
London, 1958, p. 18) remarks, 'Among the most interesting objects in this collection
_ere lhe seeds of the North Amerietn N~/umba lutl:a (Willd.) Pcrs. (No. 8517), which
lIIere grrminated by RobertBmwn in 1848, haying been sent to Slome about 1727. This
is a umarkable in~"Iance ofseeds' havinp' gcrminaIcd at a considenblc period since they
-.:n: ootlcetoo. Othcr specimens of /l;tlllmhO which, like the prco:ding, were in the
Deputment of BonUlY, were germinated b)' Bro"," between 18.0 2nd 1855; these may
Iu~ represented the same No. or possibly No. Suo, acquired by Sloane from Petiver,
- ' bentt coIJcetcd before 1718, rhe da.te of Peffi"er's death. No. 8517 is no longer
~ed in the collection, butspecirn~ of No. 81 10 sritI remain.' It is to be hoped
dill O(IX at least of lhese last can Ix induced to germinate after 1968, so amfirming
die fun qumer-millennium for seeds in dr)' storage.-Ed.

SEED PRf.SERV.o\TION .-L"ill LOKGE\lTY

Indian Lotus, in 1843-55. In 1925, 1. Ohga visired London and made

furmcr tests with these same collections of sceds. Thcy had lost their
power lO germinate SOme time between 1855 and 1926. Ohg.t, howe\'er,
succeeded in germinating seeds of the same species that had been collected
in a layer of peat in a narurally drained lake-bed in Manchuria (Ohga,
1C)23, 1926), see Plate 1('). From the information he was able to obtain
about the site in which these old seeds were found, Ohg:t thought them
to be 'at least fOUI hundred rears old' (Exell, 193J). Libby (195I), using
some of the original lot of Ohga seeds which were still viable at that time,
tested them with the radiocarbun technique and estimated that they wcre
1,-1- :L 210 years of age. These are the oldest knO\\"TI \iable seeds, su
Plate 1 (d). Libby reported, later (195-1-), that Ohg.t found three viable
LotuS seeds associated \\;th the remains of a canoe discovered about
t .....enty feet below the surface of a lake in Kemigawa, Japan, in 1951. The
average age of the wood found in this canoe was 3,075 180 years, but
this of course does not necessarily indicate the age of me seeds. There
is, however, no doubt that the seed of Nelumbo nucijera is capable of
germinating after long periods of storage in a natural condition.
It should be noted that the old Lotus seeds did not germinate without
some treatment to make the hard coat permeable to water. This was done
by concentrated sulphuric acid ueatmenr, or by the simple expedient of
filing the coat. After such coat treatment the relative growth of the seedlings from the old seeds was greater than that from fresh seeds of the same
species (Ohga, 1926b) as shown in Plate I(a). Apparently normal plants
from these old seeds have been grO\vn in many parts of the world (cf.
preceding page and Plate I(d). Reports c1aimjng that they are 50,000
years old, ho.....ever, are apparently nat valid (Anon., 1951).
The records given above of longevity of old seeds in herbaria and
cupboards include seeds wim impermeable coats which are especially
characteristic of the family Lcguminosae. Because the coats must be made
permeable to water before germination can proceed, it is vcry likely that
some of the tests were invalid, for not all of the workers appeared to have
been aware of this necessity. Much later work under controlled storage
conditions has pointed to thc efficacy of the sealing process to maintain
the moisture content fayourable for retaining viability. It is as though each
seed of great natural longevity had been individually sealed, so that the
species could be ensured of distribution in time. The importance of coat
characters in seed longevity has been rewo'llized from the t:lrliest reports
and was emphasized by Crocker (1909) as early as 1909.
EGYPTIA,'l Mm.u,fY WHEAT

The impermeable coat characteristic of long-lived seeds would seem

to discredit the perennial reports of the viability of seeds from Egyptian
tombs. For the most part these claims haye been made for wheat seeds,

RECORDS OF OLD SEIDS

though pea seeds of this ancient vintage ha\-e also come in for a share of
anention. An article in the Gardmers' Chro1/icle under the date of
Saturday, No\-ember 1I, 1843 (Anon., 1843), reported the germination of
one of tweh'e wheat seeds taken from an alabaster vase in a tomb not
visited by man for 3,000 rears. The resulting plant grew to maturity and
produced twenty-seven grains which, in turn, germinated and produced
\;gorous plants. M. F. Tupper, who germinated the seed, remarked in
conclusion (Anon., 1843, p. 787), 'If, and I see no reason to disbelieve it,
this plant of wheat be indeed the product of a grain preserved since the
time of the Pharaohs, we moderns mar, within a little rear, eat bread
ID:lde of Cam which Joseph might ha\-e reasonably thought to StOre in his
g:r:marics, and almost literally snatch a meal from the kneading-troughs of
departing Israel.'
By far the ID:ljority of reporters on seeds from tombs, however, take
the view that the seeds could not possibly be viable or that a hoax of some
kind is involved (Nicholson, 1932; Anon., 1933; Cifferi, 1942; Bunker,
1946). A description of a 'mummy' pea, so-called for no other reason than
that someone years before had claimed that the original pea of this
'-:ulery had been found in an Egyptian mummy, has been given by O. A.
\\bite (1946), who discredirs the story. Examinations of the aetllal seeds
removed from tombs ha\'e also been made. Luthra (1936) found some of
the wheat grains completelr carbonized while still retaining their original
s!upe. When moistened, these grains changed to a powdery black ash.
Barton-Wright et af. (19#) found that barley from King Tutankhamen's
tomb was extensively carbonized, but the germ with its scutellum and
embryo components was still intact. However, the old grain had apparently
lost a considerable amount of weight and density as compared to that of
fresh English barley. Also, microbiological assay revealed that riboflavin
and nicotinic acid were still present in the old barley, together with an
increased acidity. The authors thought it was possible that preservation of
the barley was helped by the oxygen-free aonosphere created by its own
~tion and the uptake of oxygen by other products stored \\:ith it.
_'lJlhing ....<IS said about the germination of these seeds. Aberg (1950)
described the taxonomic characters of samples of barley and wheat about
5.'XIO rears old, taken from the Saqqara Pyramid. An article in Nature
:\non., 1934) reviews some of the reports on the viability of such seeds
.md says that popular belief in the viability of wheat grains which have
t!rttn interred in ancient tombs, sometimes thousands of rears old, has
e\'erelr shaken by morphological and physiolO'oical tests on genuine
. my wheat, and also by bringing into question the authenticity of
c;o....called specimens.
~iocarbon dating has reyealed that some wheat and barler grain
Egypt is 6,391 1. ISo years old (Libby, 1951), but no claim is made
~~e \i.ability of such seeds.
1... ~ite of all the evidence [0 the contrarr, people will continue to be

SEW PRESERVATION AND

LO~{'nrITI

interested in, and some of them will rontlnue to believe in, the viability
of seeds from ancient tombs, though it is likely that any such viable
seeds were carried in with modern packing or deliberately placed there as

a hoax.
Much more will be said in the later pages of this book about the lifespan of seeds with permeable coats. It ",ill be seen that it is possible to
extend the life of the seed by providing it with the proper storage conditions. However, we have no evidence at present that wheat seed will
remain viable for more than tweory-five to thirty years.
For the most part, earl) records of aged seeds were obrnined from
herbaria or storage cupbo:uds. These were from seeds which just happened
to be available for testing. One of the earliest experimental designs to test
the effect of various storage conditions on seed viability was that described
by van Tieghem & Bonnier (1882). They stored some leguminous, castor
oil and flax seeds in tubes in air or carbon dioxide, as well as in open
tubcs-all at room temperature. After two years, the seeds stored in open
containers had decreased little or much in germination, depending on the
species, while those scaled in air or carbon dioxide germinated less
rodily or had lost viability completely. In the light of furure, more
accurately controlled tests, these data mean little because of the lack of
water-coorent measurements. All of the seeds exposed to air had gained
weight at the end of two years, while those in sealed containers had not.
This perhaps indicated a high moisture amtent at thc beginning of the
storage period-a condition which would bring about rapid dcterioration
in sealed storage.
Kondo (1926) recognized the importance of yariety and of c1im.1tie
conditions on the keeping quality of seeds. He stored seeds of Japanese
cultivated plants in sacks in the laboramry, and found their life duration
very short-much shorter than was recorded for seeds of Australia,
Europe, and America. He believed this to be due to the moist, hot summer
climate of Japan.
In order to investigate the effect of climatic conditions, Duvel (ll)O,lJ
stored seeds of somc common cultivated plants in laboratories in Puerro
Rico, and in different states in the United States of America (Florida,
Alabama, Louisiana, Indiana, Ncw Hampshire, and 1\'llchigan). From
these studies he concluded that precipitation is a factor of much greater
importance than temperature, Immature seeds and seeds harvested in wet
weather did not rctain viability as long as mature seeds and seeds ripened
in clry wcather. Thc life-span varied with the family, genus, and species,
Under good storage conditions the life-span could be lengthened, bur
1lC\'er, in his opinion, for centuries.

"

LlFE-SPAt'l OF SEEDS BURIED IN SOIL

IT is, perhaps, nor surprising that seeds which are dry and, as a consequence, have their metabolic processes greatly reduced, are still capable of
germination after rears of storage. The extended life-span of some species

in the soil, however, is more difficult to explain. There are many reports of
seeds germinating after ten or more years in the soil. These reports have
been based, for the most part, on the appearance of plants not common to
the vicinity-usually on newly excavated or ploughed soil, or on bombed
sites following a war. Numerous investigators have been cOllcerned with
the viability of seeds buried in the soil for long periods of time. Weed
seeds, especially, have been the subject of many studies, because of the
difficulty of eradicating all of the plants from cultivated gardens. Turrill
(1957) statcd that it has been proved at RothaffiSted that soil under
pastures contained buried and viable seeds of arable weeds after periods
of as much as 300 rears in one area and of thirry to forty years in
Others.
While much of the information about the \;ability of buried seeds has
nor permined the actual determination of the age of the seeds, some
controlled experiments have been conducted. Perhaps the earliest of these
was one started in 1879 by BeaJ (Darlington., 1951). Seeds ofrwcnty different wild species "ere mixed ,,~th sand and buried in uncorked pint bonles,
the mouths of which were tilted downwards to prevent filling ,,~th water.
Burial W<lS in sandy soil approximately 18 in. below the surface. A summary
of the results obtaincd afrer five (0 seventy rears is shown in Table Il.
It will be noted mat some of the seeds did nor withstand burial for as much
as fixe years. Again, seeds of eleven species "ere still alive after twenty
years. Mter forty years' burial, seeds of eight species, namely Amaroll1lms

'(lrofiexus, Ambrosia datior, Brassica nigra, Lepidium virginicul/l, Omothera

Mnis, Planfago major, Portulaca oleracea, and Rumex crisplts aU produced
seedlings, while after seventy rears only Oenotltera bietlllis and Rumex
uispus were capable of germination. Testing was done by transferring the
contents of the storage bottle to a 'flat' box of sterilized soil, which was
pbced in the greenhouse. Seedlings were identified after they had grown
sufficiendr to make this possible. It will be noted that seedlings of all
ri:able species were not obt:lincd at eaeh testing. Also, Verbascum b/affarja
:md Sible nOl:tif!oTIl, which were not included in the original Storage, had
~~ after fifty or SL\':ty years. The table gives a record of all the tests

TABLE II
Rf.SULTS OF ALL TESTS TO I)ATE IN m::AL'S BURII:J) SEW E.XPJ;:RIMENT

(from Darlington, 1951)

Name of species rested
AlI/r/ri:mtIIllJ rrlrojlulls

Ambrosia do/ior
JJrassiCfJ nigra
BrOil/us ueali/IliS
IJl/rSIl
~

bursa-pas/oris

Erccilti/Cs lIiaacifolia
ChlllJlQcsycc I//(/ell/%

51h

Pol)'l:olllllIJ hydropiper

Portl/laca a/noaa
RIIIIIC.f crispm
CI/(/f:IO(/,[OII luttKfm

Als"'" mtdia
Trif{)/iulIJ repem
VtrbosCl/1/I I!lapsu!
(VerboulIIlI biOi/aria)
(Siltllt lIocliflora)

35 1h

401h
}'r.

190 ..

,"'"

)"r.

19q

IIJZo

+
0

+0

0
0

".

15 1h

20th

25 1h

)'r.

)'r.

1'889

y'.
1894

"99

+
0

+
0

+
0

0
0

+0
0

Lepitlil/III t,jrgilliculII

tlgrostemllltl gilhago
tllI/brlllis colliia
,HI/IVN rOll/llilifolili
OctJO/huII bit/wis
Plrm/ago major

30lh

loth

yr.
188..

0
T
T

+0
0
0

+
+
+0
+

.,.
0
0
0
0

+0
+0
+0
+
+i
+
0

+0

+0

+
0

0
0

0
0

0
0

+0
+0
+
+
+
+
+
+
+0
+

+0
0

+0
+I
+0
+0
+

0
+0
+0

+
+
+
+0
0

yr.

+0
0
0

+0
0
0
0

I
0

+
.;0
0

+0

0
0
0

+0
+0

+0

0
0

0
0
0
0
0

+0

+
0

+0

0
0

0
0
0

50lh

y'.
1930
0
0

0
0
0
0
0
0
0
0

+0

+0

60th

yr.

70th
yr.

1940

1950

0
0
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
0
0
0
0
0

+0

+0

0
0

0
0

+
0

+
0

+
0

0
0
0

0
0
0

0
0

+
+

~
~

0
Z

~
~

"<

UFE-SPAN Of SEEDS BURIED IN SOIL

Up to 1950 inclusive, and indicates the number of times each species has
produced at least one seedling.
A similar experiment, but one which was terminated after thirty-nine
rears, was a project of the Seed Testing Laboratory of the United States
Department of Agriculture, and was started by Duvel in 1902. The tln.a1
results of this experiment have now been given by E. H. Toole & Brown
(1946). Seeds of IOj species representing borb "ild and cultivated plants
were buried in thirty-two sets in sterile soil in Bower pots with porous
clay covers at depths of 8, 22, and 42 in. Germination tests have been
reported after burial for I, 3. 6, JO, 16,20. and 39 years. Of the JO"] species
buried in H)02, 71 germinated after one year, 6J after three years, 68 after
six years, 68 aftcr ten years, 51 aftcr sixteen years, 51 after twenty years,
44 after thirty years, and 36 aftcr tlUrrr-nine years. There "'-as a general
tcndency to lowcr germination of seeds from the 8-io. depth than from
the 22-in. depth, and to the highest germination from the 42-in. depth.
The sixteen species, representing tcn plant families, having the highest
germination after thirty-nine years (more than IS per cent from at least one
depth) wcre: AbutilOIl theophrasti, Ambrolia orumiliijiJlia, CQlIVOlvulrtl
~pium.

Datura stramonium, Ipomoea locullosa" Lespedeza i11termcdia,

Sicotiana tabaCllm, Genot/ura bicfI1ris. OnopordulII acamhillm, Phytolacca
tzmaicana, Potcn/illa tlorvegica, Rohinia pIeudoacacia, Rudbeckia hir/a,
Sqbnum nigram, Trifolium pratemc, and Verbasctllll thaps"s. Of thc
orenty other species that showed some life after thirty-nine yeaTS,
Oghteen exhibited not more than 6 per cent gennination from any storage
depth.. ~lost of the seeds of cultivated plants died after onc rear in the
soil, but N;cotiana tahacum and Trifolium prtrtcnse survived for thirty-nine
~-ears. Seeds of wild plants, on the other hand, werc more resistant to
bwial than werc those of cultivated plants.
A buried seed experimcnt set up to last for forty-six years was started
10 1932 by Goss, using tweh'c weeds of California (Goss, 1939). Very low
~ation obl:ained in 1933 and 1934 '\\--as attributed to failure to supply
special germination requirements of the seeds, including treatment of
!:lttmeable coats. Many of the seeds were dead by 1938. No later report
. .!\-ailable so far as the present author is awarc. Cultivated white rice
ht'n buricd in the soil at thc depth of ordinary ploughing loses its
ubilitr during the first winter in California (Goss & Brown, 1939).
Ic:fun and California red rices behave in a similar way, though they are
~dv more resistant to burial. In general, thc seed remained alivc
-;er' in the irrigated than in the non-irrigated plots. Under dry storage
the soil temperature conditions cAisting in California, all of the red
ntt'5 ttsted showed good viability after tbIee wintcTS. This experiment
terminated after ten years, when some germination was still obtained
Conrofthe rice lots stored in irrigated soil in Arkansas (Goss & Brown,

Ani & Kataoka (J956) found a seasonal nriation in the viable seed
9

SEED PRESf.RV.o\TION A.''D LONGl."1'ln-

population of Alopecurus aequalis, a dominant weed in fields of wheat and

barley, and in its vertical distribution in the soil. The density of the y-iable
seed populations in top-soils decreased daily after the field had been
drained in autumn, the decrease v:lr)ing widely with the depth of the soil
layer and with tillage.
E. O. Brown & Porter (1942) buried weed seeds in the soil at different
depths and tested their germination after three years. The viability of
seeds of lLpidium draha L., Lepidill1n repau L., and lIynulIopllysa
pubescens C.A.Mey, deteriorated rapidly when buried in soil at a depth of
4 to 6 in. Seeds of Eupllorllia escula L. kept well at 6- and IS-in. depths.
eenlaurea ca!cilrapa L. and C. solslilialis L. had a short life-span in the
soil, declining from 90 to 10 per cent germination within three years. Seeds
of C. repens Boiss. were less subject to deterioration. Donnancy in
Solanum seeds present at the beginning of the test disappeared after three
years of burial, but the actual germination of seeds of S. elaeaglliftlilll1l
uv. was grcady impaired by three years' burial at IS in. Ninery-eight
per cent germination of S. carolinmse L. seeds was obtained after burial
for three years at a depth of 4 to 6 in., but very few survived burial at 16 to
18 in. for the same period.
One of the remarkable things about the longevity of weed seeds in the
soil is dut most of them do not have impermeable coats and hence absorb
water immediately upon exposure to moist soil. That seeds which
have imbibed their full complement of water should remain viable over
such a long period is noteworthy. Furthermore, most of these seeds
can scarcely be considered dormant, though some of them may de\-c!op
secondary dormancy. That they do not develop a 'deep' dormancy, when
buried in the soil, is demonstrated by the fact that when the soil in an
ordinary garden plot, for example, is disturbed by culti,ration, many
weed seedlings appear even if care has been taken to remove all such plants
for several preceding years. Exposure to light, alternating temperature,
mechanical disturbance, or some other unknown. factor may SCTve as a
stimulus to bring about germination of these buried viable seeds. Furthermore, it has been shown by Kjaer (1948) that some seeds actually remain
viable longer in the soil than in dry storage. His report includes tests of up
to a ten-year period at the end of which nine of the different seeds mat
germinated after burial failed to germinate after dry storage. Out of the
seventeen stored, only one species, Gcrtlllit/111 disseClu1l1, survived the dry
storage but failed to germinate after burial in soil. The other seven species
("tiled to germinate after ren years regardless of the storage condition.
OthCT evidence of the superiority of moist soil over dry storage at room
temperature is to be found in \'arious reports (Crocker, 19-+8, p. 40).
Seeds of the Western binerweed, Aclinca odorala, were stored in the
open, fully e.'(poscd to weather conditions, and in the laboratory in Te.'<as
(Reynolds ~t al., 1938). Both lots of seeds had a viability of 72 per cent
at the beginning of the test. Both lots lost their germinating power with
10

1.If'E-SP.\o~

OF SEEDS UUKlfD IN SOIL

increasing age, but the seeds oUlside deteriorated more rapidlr than those
in the laboratory. All of me laner were, however, dead after fivc rears of
storagc.
MOIsr STORAGE OF SEEDS all

Amaral1lhUJ refrojle:rus L.

Vlhen seeds mke up watcr preparatory to germination, their metabolic

activities become greatly accelerated. It is evident mat seeds which have
imbibed water could not remain viable for very long in the soil unless
definite curtailment of these activities took place. In an effort to obtain
more definite information on the effects of burial, freshly harvested seeds
of the pigweed, Amaranlhus retrojfex/ls L., were used (Banon. 1945h).
Seeds of this plant when freshly harvested show a degree of dormancy
which varies from year to year and, with seeds collccted in "'3rious
localities, in any onc rear. The 'dormancy' is expressed in lhe requirement
of a high temperature for germination. The best temperature for germination of freshly harvested seeds was 35C., at which 86 per cent of seedlings
,\-ere obtained from one lot and 73 per cent from another lot immediately
after collection. Furthermore, the tcmperature \\'3S quite specific at that
time. Some germination occurred at 25 and 30C., but the percentages
here did not exceed 18 in eimer lot. Only occasional seedlings appeared
o\"er the rather wide range of other temperatures tried.
When these seeds were stored dry after harvesting, a gradual change
in their capacity for germination took place, so that after tWO or three
months they germinated over a \\ide range of temperatures including
20C. If, however, they were placed on a moist medium at 20C. immediately following harvesting, this after-ripening did not proceed but, rather,
primary dormancy was maintained or a secondary dormancy developed.
This resulted in a different pattern of germination behaviour.
Careful records of the germination on moist glass-wool at 20C. of
four lots of Amara1llhus retrofleJ:uJ seeds ha\-e been kept for eight years
(Barlon, 1945b). All four lots were similar in germination beha\iour. A
summary of the germination of nyo different coUections made in 1942 is
shown in Table III. These seeds have exhibited a periodicity in germination which has been remarkably uniform for the four lots" It may be noted
that the data in collection A are based on twenty-two lors of 2,500 seeds
e::tch (55,000 seeds), whereas those in conection B are based on eighteen
lots of2,500 seeds each (45,000 seeds). Germ1n.1.tion behaviour ofindividual
Ims of 2,500 seeds was recorded, but ail lots of a single collection have been
combined in the figures shown in Table III. The number of seedlings
appearing by the end of each nm-month period during each of eight years
is given. Collections were made in October, so that the first n\"o months
represent October and Tovember and the last two months represent
_-\.ugust and September. Within a period of two months after the seeds had
been moistened and placed at 20C., a toW of 186 seedlings had been

"

SUD PRESERVATION A.'\'D LQXGEVfn'

produced from seeds of collection A and 378 from those of collection B. A

strong gennination-pulse was noted for the tenth to twelfth months of
storage during the first and succeeding years up to eight. This periodicity
of germination is apparently indcpendent of e.xternal conditions, for it took
place under nearly constant conditions. This indicates a v'lI')'ing degree of
the primar)' dormancy or the induced secondary dormancy of the
individual seeds of the original lot. Moist AmQrQ1lthus seeds, held:without
germination at 20C., could be induced to genninate at that same temperature by rubbing, by drying for from three hours to three days, or by
exposure to 3SC. for twelve to twenty-four hours. Germination also
proceeded immediately aftcr removal to higher constant or alternating
temperatures.

TABLE III
Amaral/rIms rerroflexlls,

19.f2 CROP. GER.\-UNATION AFrER VARIOUS PERIODS

DURING UCH OF 8 YEARS ON MOIST GLASS-WOOL AT 20C.

No. of seedlings produced each z-monthly period dueing

Yrs. of

Collection

mo'"

the year

storage {Oct.- (J)cc.- (F~b.- (Apr.No\.) Jan.) Mar.) May)

A
55,000

"""

Total

,im

8
Total

"
7"1

47
83'

IZ,201
12,8 14

:::37

'47

'7

'0'

5
o

151

'5

"
,

"'0
66,

.,

,..57

12,763

.8,

3,155

1,132

1.308

7-H

43'
5,638
4. 829

16,507
49J
6,C4-7

135

'3

5
3,80z

1,163

39,82 9

2,994

o
"
337

18,zo8
5,454

6,659

7'
5

u,66

13,70 3

26,575

8,
z5

3-1

454

1,223
4,227
73 8

1,568

378
8,

60

.06

&p,.) Tow

455

'

"

5"

(Aug.-

47
17 1
74

899

.".

'0

u=Jul.)

3J

".

, "

",

".

1,732

'73

".".

435

7.010

973

6,324

45'~

:::,358

75

253

2,381
28,034

9 15

7,66t

,,8

3,3 24
42,40 1

Gaseous exchange of Amaranrhus retrojkrus seeds, measured at

intervals of from 0 to 1)01 days of moist storage at 2Q 0 C., showed at least
a ten-fold reduction in respiration. The beginning ofthis reduction became
apparent very early (after two days), and was definite after eight days in
moist storage. The decreased respiratory rotc is probably an indication of
>Z

LII'E-SP.o\"- OF SEWS Bl.llIID IN SOIl..

the reduced metabolic activity of seeds in the soil which have imbibed
water, though it is believed that further reduction from that measured
here would have to take place to permit these seeds to li,-e for as long as
fortylear5. The present experiments were discuntinued after eight rears.
Toole & Toole (1954) found that seeds of se,'en lettuce varieties which
had imbibed their full complement of water rem,!ined viable on moist
filter-paper for J05 days, whereas dry seeds in a humid aunospbere were
dead after forty-two or tv..enty-{mc days, depending on the variety.
Storage was at 30C., a temperature known to maintain dormancy of moist
lettuce seeds.
Additional studies of respiration as well as of other processes taking
place within the seed are needed for an explanation of the physiological
responses of buried seeds.
In thc case of seeds with impermeable coats, it is obvious that their
germination will proceed in soil 'storage' as soon as micro-organisms or
mechanical abrasion makes the coats permeable to water and sufficient
moisture is available. The severallarcrs of the seed-coat of Ntlumho are
shown in Plate I(h), with the parts which prevent water absorption
indicated. In the old seeds that bad been in the soil for hundreds of years,
the epidermis and mueh of the outer ends of the palisade cells had disappeared, but most of those old seeds which were recovered from the soil
still required coat treatment for germination (Ohga, 1926). This probably
indicates simply that such old seeds as became penneable over the years
germinated, but, because they were buried at such 6rreat depths, the
resultant seedlings failed to survive.

'3

1II

MOISTURE EFFEcrS
MAt....' Y factors, such as moisrure, temperature, gaseous exchange, seed-oJ3t
character, maturity, microflora and insect infestation, may determine the
longevity of seeds under natural or controlled stOrage.
It has been well established by many investigators that the moisture
content of seeds is of the utmost importance as a factor in the determina-

tion of their longevity. This applies not only to the absolute moisture
content but also to fluctuation in moisture content-especially to fluctuations around the 'critical' moisture content which varies according to the
type of seed. That temperature and gaseous exchange are of vital importance in connection with the keeping quality of seeds has also been

demonstrated repeatedly and will be discussed below. None of these

items can be considered alone, as the effect of one depends upon
others.

The importance of moisture in the storage of seeds has been recognized by the Inrernational Seed Testing Association iil. the fonnation of
a Committee on Seed ~(oisture Conrem and Seed Storage. This Committee obtains information on seed moisrure and seed storage stUdies
made in different areas of the world and serves as a dearing house for
such information (E. H. Toole, 1957). As early as J934. the appearance
of abnormal seedlings which caused difficulty in the interpreration of
germination tests was attributed to high moisture content of the seeds
at some time after harvest (E. Brown et 01., 1934).
In view of these facts, it becomes highly desirable to ascertain the
actual amounts of water which will be absorbed by seeds in different
localities, i.e. under different conditions of humidity and temperature.
Such knowledge would make it possible to determine whether air-dry
seeds, in any particular region, could be stored with safety. Data on
moisture contents of six varieties of seeds, stored :J.t four different temper:lfures and three different relative humidities, have been presented by
Barton (1941). Correlations with the germination capacity under all of
these conditions have been m.:t.de. As the resultS of these tests have a
direct bearing on keeping quality of seeds in general, they will be presented here in some detail.
Seeds used were oflerruce (Lactuca sativa L.), onion (Allium upo L.),
tomato (LycopersiCllm tScu/rotum Mill.), flax (Linum usitatissnmw: L.), peanUl (Arachis hypogata L.), and long-leafed pine (Pinus pa/lfstris Mill.).

'+

TAULE LV
MOISTURE CONTENTS OF SEJ;.DS STORf.D AT VAIUQUS TEMPERATURES AND RELATIVE HUMIDITIES
(MOISfURE EXPRESSf.J) AS PERCENTAGE OF DRY \VEIGHT OF SEEDS)

Per cent moisture after storage for 8, 29, 105, and 372 days
Sc~d+pcr c~n[
mOisture at time
of storage

Lettuce
6'5

Onion

35

55
76
35

10'2

55

76
Tomato

89
Flax

7. 6
Pine

8,

58

'9

83

68
9'
9. 6

10'7

108

7"
T5

lIg

13'7

12'8

IS" I

20"C.

lODe.

"5

37'

5'8
8,

6,
87

84
10'Z

10'5

12'7

10'1

13"3

19'6

15"7

9'

99

12" t

10,6

10"'

10'5

13'4-

5' J

9'

12'4
IS'2

15'4

15'1

I.l."o

15,6

136

17'3

13'7

IBS

30'5

lS'7

14'.j.

89

37'

68

59

10'0

8'5

6,
9'

'9

10

37 2

H
9. 8

S"2

,..5.8

10'2

12'3

rI"}

II'S

86
83

83
!J'O

,6'3

9'
11'8

17"1

ITS

97

93

9'

8,

106

9. 8

83

55

II"

93

15'7

I:d~

11'2

12'9

IS'S

I7'!

1+'4
17'3

13"]

76

11'413'8

8'5
13,6

85

12"2

15"7

18'5

nB

11'7
14"1

3.1

8,
86

74
99

76

68

12"6

12'0

76

68
93

9"5

10'4

T4
10"8

II'O

9"5

II'S

II"4

Lj"'.2

11"0
16'5

20"4

10"6

3.1

8"S

86

89
9'

10'5

7'
98

11'1

II'6

6,
63
67

6,

48
7'
86

55

55

35

5.1
76

77

g.o

10,8

Q'S

76

68

8,
9. 6

12'2

14"4

13"8

15'0

8'3

8,

"1
".

T5

63

12"6

!:;l'6

13'0

IZ'S

63

69
9'

10'3

78

83

T4

99

[l"6

14"6

10'9
10"9

10" 1
12'7

".8i

7'
8,
7'

6,

4. 8

8"5

T7

59
8,

f9
4

.I'

13"9

12'4

8"5

10'0

9.6

7'
88

12" 1

II'Z
16'0

n'I

85

67
9. 8
9. 6

74

77

59
9'

5'0

59

10'4-

67
87

10,8

45

9'

6,
8,

85

12"7

7'
97

12"9

5'
65

37'

10"7

10'3

9'

105

10'4

15"4

II'I

'9

10'0

97

T7

74

'5'9

10"9
13'9

"5

10'2

7. 8

Jae.

'9
6 ,
85

35

76

Peanut

Soc.

Pcr cent

ReI.
hum.

13'4

j.8

.,

10"6

7'
9. 8
49

7. 6
7'

93

63
77

T7

13'1

53

5'9

II'I

6"4
Il'Z

7'
11"6

64
8,

5"5

6,

69

TI"6

II,g

7"5
IZ'3

4. 6

39
4. 6
97

5"6

87

I
~

SEED PRF.'iliRVATION At"ID LONGEVITY

Large d esiCC3tors containing saturated solutions of magnesium chloride

(MgC1 2 . 6II:O), calcium nitrate (Ca(N0 3) : . +lI 20), or sodium chloride
(NaCl), with an e'(cess of the salts, were placed in constant temperature
chambers of 5,10, 20, and 30C. These three salt solutions provided
atmospheres of approximately 35, 55, and 76 per cent relative humidities,
respectively (H.M. Spencer, 1926). The seeds were placed in muslin bags
over the solutions in the desiccators. Moisture determinations were made
oneaeh seed-lot after 8,17,29,43,15.15,232, and 372 days. by drying in
a vacuum oven at 78C. for forty-eight hours. All tests for the germination
C:l.pacities ofthe seeds were made in ovens at controlled temperatures wmch
had previously been detcnnined as favourable.
Results of moisture determinations made 8, 29, 105, and 372 days
after storage under special conditions are shown in Table IV. Tests made
after eight and seventeen days on lots representing all of the different
conditions showed fluctuations which indicated that the seeds had not
reached an equilibrium with the moisture in the storage chamber. Tests
made as long as twenty-nine days after storage showed that an equilibrium
was being established by that time, and changes in moisture content of
individual samples observed on subsequent dates were doubtless due in
great part to the error of sampling.
It was noted that the seeds showed differential water absorption
according to the species, and thar the amount of moisture taken up under
all conditions by each seed type always appeared in the same relative
position. This is shown in Fig. 2, where the moisture determinations
Ill:J.de forty-three days after storage arc used. In the order of increasing
water-absorption capacity, the seeds wcre of peanut, lettuce, flax, pine,
tomato, and onion. This order persisted regardless of storage temperaturc
and atmospheric humidity.
The chemical composition of the seeds may account, at least in part,
for this variation in the amount of moisture absorbed by different seeds
under identical conditions. Peanur, lax, lerruce., and pine seeds, which
contain large amounts of oil, have low retention capacities. On the other
hand, tomato and onion seeds, both of which contain larger amounts of
protein and smaller amounts of oil, occupy intermediate and high positions as regards ability to absorb moisture. EVi':l.rt (1896) stated that the
resistant power of a seed to desiccation depended partly on the nature
and thickness of the coat and partly on the form in which the reserve food
was stored. Other conditions being similar, albuminous seeds were least
resistant to desiccation, oily seeds next, and starchy seeds most resistant.
The present tests confirm these statements for albuminous and oily seeds.
The types of seed-coats, also, doubtless playa role.
It should be pointed out that, at the beginning of the tests, seeds of
the same species possessed the same viabilit), in all storage conditions;
but, as the storage period lengthened, loss of viability progressed much
more rapidly under conditions of high humidit), and temperature. This

,6

~IOlSTURE

EFFECfS

'0

"
I.
Onion

"
Tomato

"

10

'"
~
~

...

I.

...Z
'"
0

..'"

Lettuce
Peo.nu1

55%

Pine
Fin

I'

"

10

Onloo
Tomato

Pine
Flax

Lettuce
Peonu1

'0

10

'0

LeHuce
Peonllt

'0

TEMPERATURE "C.
FIG. z_~Grophs indicating moisture content of seeds after fort)--thrce days of stonge at
nrious tcmpe1'2tlll"CS and at relative humidities of i6, 55, and 35 per cent.

means, then, that moisture determinations were made with increasing

proportions of non-viable seeds_ That the state of viability was withouJ

'7

SEED PRESERVATION

_"-''\;1)

LOXGE\lTI'

effect on the actual amOunt of water absorbed, was demonstrated by the

failure of the moisture: COntent [0 change in relation to decreased germination capacity. This may be seen from a comparison of Tables IV
and V. Other workers have also shown that seed viability does not affect
the total water absorption. Heinrich (1913) weighed seeds of Lclium
puettne after 0 to 384 hours in a moist room at 20"C. and found that
li\>ing seeds contained more water than dead ones up to twcory-four hours,
aflcr which they were approximately the same in this respect. Atkins
(1909) found that seeds, whether living or dead, take up the same quantity
of water in their initial stages and that there is no difference until germination begins, and Simpson & Miner (1944) found essentially the same
moisture contents in living and dead seeds.
With exposure to a relative humidity of only 35 pcr cent, seeds took
up approximately the same amount of water at 5" and IO"e (Darton,
1941), but in evcry case they remained drier at the higher temperatures of
20 and 30C. (Fig. 2). At 55 and 76 per cent relative humidities, however,
the peak of moisture absorption was at IQ0c. and the least absorption
was generally at 30C. There was no indication that equilibrium was
reached more quickly at 30"C. than at any of the other temperatures
tried. Boswell eJ al. (1940) stored seeds of bean, sweet corn, beet, cabbage,
carrot, onion, spinach, tomato, and Spanish peanut at 50" and 80F. at
relative humidities of 78 to 81 per cent, 66 per cent, and 44 to 5' per cent,
and reponed that at a given relative humidity the seeds developed slightly
higher moisture contents at 50" than at 80F. These authors stated that
this could not be explained by the physical characteristics of the air, such
as differences in vapour pressure, weight of water in the air, or vapour
deficit.
It has been reported that seeds of blue lupine and Austrian winter
field pea absorb moisture more rapidly at 35" than at 20"C. (McKee &
Musil, 1948). According to A. 1- M. Smith & Gane (1939), the rate at
which moisture is taken up or lost by a eenain kind of seed depends upon
the relative humiditv of die air and the thickness of the mass of seeds.
They studied the h);groscopic properties of Chewing's fescue grass seeds
after drying. The water COntent of wheat grain at a given relative humidity
is less at 25C. than at 10"C. (Gme, 1941). Gane (1948) also found a
difference in water content between different seeds exposed to the same
relative humidity, and a decrease in water content with increased temperature at a constant relative humidity.
D. A. Coleman & Fcllows (1925) determined moisture contents of
cereal grains and flax seeds exposed to 15, 30, 4-5, 60. 75, go, and 100
per cent relative humidity at 25"C., but no other temperature ,,-as used.
Darton (1941) showed that there were significantly higher moisture
contents of seeds at a given relative humidity at 10"e. than at 30"C., in
spite of thc greater amount of moisture available in the atmosphere at
the higher temperature. Less moisture was absorbed at 20"C. than at

18

TAllLE V
GERMIKATION PERCE......'TAGES OBTAlf\.'ED FROM SEEDS STORED AT VARIOUS TEM.PERA'ruRES AND RELATIVE IIUMIDITIF.S

"'oJ

Ptr tenl gcrmin:tion .rter 5101'llge for 8, 43,

- p:r cerll Per

gnminallon cent
ar time of
SIQrotgc

Lettuce

6)

Onion

"
~

ReI.
hum.

43

JS

59

jg

'5
7'

S'
S'
S'

'7

10C.

's' ')'
7'
78

"

97

6,

II" 3'"

JS

0.,

57

3'
55

55

88

9'

93

97

,6

'J

53

88

JS
55

8,

9'

88

97

,6

97

94

JS

97

54

JS

'00

'00
'00
'00

49

'00

'"

~ 'S,

""

-59

Peanut

43

66

55

7'

9'

I'inc

37~

" li '7 !~ t:
il " " "
",. " " " ,.
0'
""80 "
1;"
7' I',
0'
"
,.
,8
,8
!1
7'
" ~ " Ii " ,."
,8
7'
- 8,87
~
7'
7' 7'
""
it
87
S'
" ""
,8
"
- ,.~ ,8
" il ''""
66

Tomato
93

Floor

5C.

'00
'00
'00

'00
'00

"3'
33

Ii"

,'I
78

,6
47

"

'3'
7'
57

53

""

37'

",8 ""'" S',

n ,8 "" ",,
" "

.,,

3'
,3

53

",

7'

95
93

~~

,6

"87

"
0'

OJ

7'

il

-" -,6
96
-

i~

"

JS
)

'00

Indicates that no

tCSI

'3'

~~

7S

.,

20

'S'

37'

')

~~

'5o,1J2 anti 373 days

"

'00

:i

was made.

63

I;
83

59

3'
0

, "3,

~J

88

8:

7J

95

"it ",8

')(1

IS

oJ

I;

"

'~"

"

68

8,

l'", !~

~6

3'

88
87

9'

'00
'00

60

OS

'00

98
,8

,,

"",

3'

'00
'00

,8
98

"

",

, ii,

, ~

83

79

~;

" ;1 ,.ii ", ", ,.,

86

8,

'00
'00

37'

))

-'7
:1
",6 , ", 98
98

')'

54

94

'00

'"

,
,
)!
6
"" ", , ", ,
'7
""3 ",, '3,, '5,,
,6

3'

8,

.,

30'C.

SEED PRf.5I.RVATION ,\,',,'1) I,ONGEITY

100

80
60
40
TomCito

P'"''
Fhu

20

Onioo

7.~

Pille

Lettuce

0
100

Paollul

Z
0

"'"
z

FloJ
Tornillo

'0

'"w

60

....z
w

..,

40

'"w

20

..

LilluC8
,,~

....Onlon
Pine

100

Peonut

TOlllato

60
60
40
Llttlle.

Pi,.

3~''4

20

Onion

10

20
TEMPERATURE C.

FIG. 3---Graphs indicating germination after stor.lge for 150 days at nrious temperatures
and rclall.e humidities.

IOe., but more than at 30C. Apprm:lmate1y the same amount was taken
up at S and mOe. at 35 per cent rcbti,c humidity but, as the relative
20

MOISTURE EFFECTS

humidity was increased to 55 and to 76 per cent, seeds absorbed more

water at IO"C. than at S"c. The explanation of these phenomena must
be sought, not only in the physical conditions of the atmosphere surrounding the seeds but also in the physical conditions of the seeds themselves. The actual determination of the vapour pressures of both seeds
and annosphere at the different temperatures used. and the relations of
these figures to each other, may provide a basis for explanation of the
beha,;our.
Yiability tests made by Banon (If}-l.I) after 150 days of storage are
shown in Fig. 3- A comparison of these graphs \\ith those in Fig. 2
reveals that the actual amount of water absorbed by the seed is not
clirect1r related to its loss of vmbiliry under any gi,"en set of conditions.
The three seed types exhibiting greatest retention of viability under all
conditions were peanut, flax, and tomato. On the basis of the actual
amount of moisture absorbed these seeds occupy low, intermediate, anu
high positions, respectively (Fig. 2). Similarly, lettuce. pine, anu onion
seeds, which lose viability mOSt readily under all conditions, have low,
intermediate, and high moisture COntents, respectively. Al<;o, deterioration does not appear to depend upon composition of the seed.

TABLE VI

s.m

ESTL\1.-\TID M.\..UMm1
SEED .1\10ISTlJRE COl\TE.."lS FOR STOR.<\GE
FOR O:xE Yf.,\R AT DIF'FERE.'T MF..-\t.'\; TEMPERATURI.S Of'

STORAGE (approximate guide only)

Kind
of seW

:\h:cimum gfe seed moisture rontcnt for :I,\'erage

temperature of storage
indicated
4S- IOc.

21 cC.

'5
'5

u
u
u

Kind
of seed

26;cC.
4O'";o"F. 7"1" SecF.
per emf pet" ernt per cent
Bean, Kidney

Dean. Lima
Ike,
Cabbage
Carrot
Celery
Corn, Sweet
Cucumber
Lettuce

"

'3
'3

"

7
9
9

'0

>0

Spcci21 precautions

8
8

l\laximum safe sttd m0isture contCllt fOf an~r2ge

temperature of srarage
indicatl

..j.;_locC. 21C.

26;CC.
c
joT.
Se"F.
4a-so F.
per cent per cent per rent

m~

OnIDn

"u

"8

Pea. Garden
'5
'3
Peanut
5
(shelled)
6
7
5
Pepper
>0
7
9
Spinach
8
u
'3
Tomato
8
u
'3
Turnip
8
5
'0
Watermelon
8
'0
needed wh~ removed to higher tempenture.
9

>0

6
9

3
7
9

6
7

Boswell et al. (1940) also reported that among different kinds of seed,
detcriorntion was not always correlated with the relative moisrureabsorbing capacity of the seeds.

SI.1)

PRESER\'ATION Ai\"D LONGEVITY

E. H. Toole (1939) !:>'1:atcs that the relative loss ufviability of different

kinds of seeds at a given rempcrarure and air humidity depends in part on
the moisture content and in part on the nature of the seeds. Tomaro
seeds will reum viability longer than onion seeds at a given temperature
and air humidity, but the longevity is fairly comparable at a given temperature when the moisture content of the seed is the same. On the other
hand, peanuts take up comparati.cly little moisture, but they deteriorate
rapidly at high temperaturcs even with a low moisture contenL l'vlaximum
safe seed-moisture contentS for storage for one year at different. storage
temperatures have bt..-en estimated by E. H. Toole (1lJ42) and arc giVCD
in Table VI.

MolS'I'UKf R.EL"TI~IPS OF Sons STORED 'OPEN' IN THE

LuJoRATORY

Whether 'air-dI)'> seeds contain safe moisture contents for storage

depends entirely upon the locality and rime .of year that storage is to be
accomplished.
In order to determine more cxacdy 'what fluctuations actually t:ll:e
place under ordinary :ltmospheric conditions, during the course of a ye:uat Yonkers, New York, moisture determinations were made in FebI1l:lr)',
.May, July, August, September, Nm-ember, and Det:ember 1939, :md
January 1940, of seeds of carrot (Daucuj CiJro/a. L. var. sa/rca nc.),
eggplant 'aubergine' (Solanum nu:/ongena L.), lettuce, tomaro, and lon.;leafed. pine stored open in the laboratory. The results of these tests showed
that tile moisture content of seeds in August ,,-as practicaUy double that
in Januar)' or Febnury. This applied to e~ery type of seed tested,
although there ,vas considemble variation in the actual amounts of water
contained in the different ,-a.r1etics. These differences are shown graphic:illy
in Fig. 4, where a comparison of lettuce, pine, and tomato seeds is made.
In spite of the differential 'wOlter-absorbing capacities of these seeds, aU
of them contain the ma.Umum moisture in August and are relatively dry
during the montJIS of November, December, January, and February. The
moisture contents of carrot and eggplant seeds at the various testing
periods were practically the same as for tomato seeds. Pine seeds took up
less moisture from the air, while lettuce seeds remained driest.
This tcst simply serves to emphasize the necessity of drying seeds to
:1 known moisture content before storage, as the amount of moisture
present in 'air-dry' seeds depends not only on the locality but also on the
time of year. 'Air-<lry' seeds placed in sealed containers during August,
in Yonkers, New York would COntain twice as much moisture as the same
seed-lot sealed during the winter months. In the evaluation of sealed as
opposed to open storage, r.hese facts should be considered.
The differenti:J.1 absorption of mI.ter by the different species furnisht:S
additional evidence that the 'critical' moisture content varies WitJl
22

MOlSTLRE EfFECfS

dilferent seeds. It is belie'oed that fluctuation in moisture content is most

deleterious to seed viability when it occurs around the 'critical' point.

"
10

.,
<5
w

'"
~

'"w

..,
~

~:::::::Tomoto

PIn,
....-

....-

..... L'It\IC'

FEB. 1939

MAY

JUlY AUG. SEPT,

NO\l

riG. ;f.-Gnph showing: mmsture mn!cnrs a! \'1rious limes of me

DEC.

~'ear

JAN. 1940

of seeds

'open' in me bbor:l{ory. J\loisrnre is expressed ll.~ pcrcent;tge o( dry-weight o( seeds. Duplicnes

of appro~imalcly :l. KJI1.. t':lch wcre

used (or the lests.

The imparlance of fluctuation in moisture COntent on the keeping

quality of dandelion (Taraxacum oificillole 'Ycber) seeds bas been invcstigated b)' Barton (1939h, 1953), who stored thc seeds at three temperatures:
boor-nary, 5C., and _+oc. Air-dry seeds at the time of storage cont:tined 7'9 per cent moisture. Some samples were stored with this moisture
content, while other samples of the samc seed-lot \\ere adjusted, by
storing in desiccators over calcium oxidc, to 6'2, 5'0, :lnd 3'9 per cem
moisture before storage in sealed containers. Sealing was accomplished
in twO ways. In one case, lotS of approximately 400 seeds each were placed
in small glass tubes hermetically sealed in air and in a vacuum; in the
other case, larger lors were placed in tin cans with tight-fitting lids sealed
\\;th se'Jling wa.~. Each tin an was opened and samples were taken out
at intervals for testing, after which the ans were re-sealed. Thus the
seeds remaining after each germination test hnd been subjected to fluctuation in moisture and gaseous exchange attendant upon the opening of the
can-a condition which did not exist for individual testing samples stored
in glass tubes. The mcthod of sealing pron:d important in keeping-quality
as shown by Fig. 5, which depicts the resulrs of extcnded storage for up
to sixteen years of seeds with initial moisture contents of 7'9 and 39 pCr
cent. Seeds with 7'9 per cent moisture (Fig. S(A stored in a tin can in
2)

SEED PRESERVATI01\ Ai"D LONGEVITY

the laboratory lost germination capacity completely within two rears,

while those stored in sL'aled glass tubes still gave 66 per cent germination
at the end of that time. Furthermore, I2 per cent of lhc latter germinated
after four years' storage. An eycn more striking difference in the effects of
two sealing methods is to be seen in the 5"C. storage. In tin cans a high
germination capacity was reL1.incd for four years, after which there 'was
Tather rapid deterioration up to ten rears, when only 7 per cent of the
seeds were alive. In glass tubes at S"c., on the other hand, practieally full
viability was retained for fourteen years, at which time the supply of seeds
was exhausted. Even at the more favourable storage temperature of
~4"C., the advantage of the glass tube over the tin can for keeping these
seeds is evident, though the 'break' in germination representing the
beginning of deterioration in this case was delayed lllltilafter t"'elve years
of storage.

~---"

r
L->-~=~-~~_-',--_~l-~_~_---l

12

!6

12

Ii;

YEARS OF STORAGE

FIG. 5--Graphs showing dfcet of h'mpcr-.uure and method of sr:aling on tbe "iability of
stored dandelion se.:ds II-irh an initi:>..! CQ/ltent of (A) i'9, and (D) 3"9 per ceut moistrn-e.
Germination al rime nf star-age was 1)1 pt'r =L

The life-span of dandelion seeds may be e.\:tended by drying to a 3"9

per cem moisture content before storage as shown in Fig. S(B). In this
case, deterioration in a till can in the laborarory became marked onIy after
four years of storage, but the fayourable effect of storing in glass tubes is
still evident. Similar effects are to he seen for Soc. storage, but they were
not evident for -4"C. storage up to sLueen years.
The- more rapid deterioration of seeds stored in scaled comainers
which are repeatedly openel! and re-sc..11ed, as compared with seeds in
scaled containers which are never opened from the time of storage to the
time of testing. sheds some light on the moisture relatiuns of seeds.

24

:3

'"

:3

MOISI1JRE EFFEcrs

E,;dence in the form of delayed ill-effects of opening and sealing, by

decreased initial moisture content, of the seeds of dandelion leads to the
conclusion that fluctuation in moisrure content is harmful Reference is
made again to Fig. 5 on this point. Moisture determinations. made by
drying the seeds in a ,-aeuum oven at 7SC., have shown that seeds with
a moisture content of 7'9 per cent at the beginning will gradually dry out to
about 6 per cent with repeated opening of the storage can. As far as
moisture content itself is concerned, this should be an advantage, but the
fact is that dandelion seeds 'keep' better with a constant moisture content
of 7"9 per cent than "ith a fluctuation downwards. Deterioration of seeds
with 3'9 per cent moisture is accelerated when repeated opening of the
can has brought the moisture content up to about 6 per cent as shown by
actual moisture determinations. Dandelion seeds ,\ith intermediate
initial moisture contents (6'2 and 5'0 per cent) were intenned.iate in their
responses to moisture fluctuation.
It should not be assumed from this discussion that it is always better
to seal seeds "ith high moisture content than to leave them open. If the
moisture content is excessive, there is increased injury in scaled conrainers; but within certain limits, which probably vary for each kind of
seed, fluctuation in moisturc content results in greater damage to the seed
than could nonnally be expected from constant levels of moisture content,
Subsequent to the findings just described, a test was set up designed
tD include controlled fluemation of humidity at a constant temperature of
20~C. (B:mon, 19-1-3). Desiccators ,,;th different relati,c humidities were
used as storage chambers, a constant humidity being maintained in each
desiccator by means of a saturated salt solution ,\ith an excess of the salt
(H. M. Spencer, 1926). Fluctuations in relative humidity between 35 and
55 per cent, between 35 and 76 per cent, and between 55 and 76 per cent,
at intervals of two, four, and eight weeks, were made by transferring the
bags containing the seeds from onc desiccator to another. As controls,
certain lots were left at each of the constant humidities for the entire
storage period. The sl.:cds used were of onion (AJlium cepa L.), dandelion
(Taro:cocum ojficinule Weber), eggplant (So/anum me/ongroD L.), and
roma(Q (Lycopersirom escu/mlum MilL), the first two being tricd because
the)' arc yery sensitive to ad\ersc storage conditions and the second two
because they arc relatively resistant to the samc conditions. Viability tests
were made on moist filtcr paper in ovens at controlled temperatures,
except in the case of dandelion which was germinated in the laboratory.
In general, when alternation from one humidity to another was
effected, the variation in moisture coment of the seeds tended to be the
SlIDe as was indicated by the determinations madc from consranthumidity storage. I n some cascs, especially if the alternations were made
C'ycrr two weeks, \-ariation in moisture content of the seeds w-as somewhat
lesslhan lhatindicated bythedclerminations made fromconsranl-humidity
storage. Seeds transferred from a high to a low humidity failed lO reach

2;

SEED PRF1iF.R\'ATION M\.'D I.QNGF:Vrrv

the low moisrure content expected before they were transferred again to a
higher humidity, where again they failed to develop the maximum
moisrure content to be expeered.
Onion seeus held at constant relative humidities of 35, 55, or 76 per
cent acquired moisture contents which averaged 8,6, n'4, and q.9 per
cent, respecti,-e1y, and remained at these values throughout the tcst
period, Onion seeds which were alternated between 35 and 55 per ccnt
relative humidity attained an average of 9'4 per cent moisrure after nYO
weeks at 35 per cent and 1 I per cent after two weeks at 55 per cent, so
that the average ,-ariation in moisrure content was 16 per cent CVel}' twO
weeks. Similarly, the average ,-wation when the seeds were changed from
3S to 76 or from 55 to 76 per cent moisrure every two weeks was 4'7 and
2 per cent, respectively.
Except at a constant relative humiditr of 76 per cent, neither eggplant
aubergine' nor tomato seeds showed any reduction in viability under
an}' of the storage conditions up to the end of sixty-four weeks, at which
time the experiment was terminated, These seeds arc kno,yn to be resistant to unfavourable storage conditions. A constant relative humidity
of 35 per cent imparted a moisrure content of 9 per cent to onion seeds
and permined the retention of their viability for one year at zoe., with
the percentage germinatioll only slighdy reduced at the end of that period,
At 55 per cent relative humidity, at which onion seeds came to conmin
about I I per cent of moisrure, serious deterioration occurred after twenty
weeks of storage, At 76 per cent relative humidity considerable loss in
germinative power was evident after eight weeks of storage, and, after.
twelve weeks at. this humidity, where the moisture content of the seeds
was about 15 per cent, onion seeds yielded only Z3 per cent germination
as compared with 96 per cent at the beginning of the speciaL storage
period.
These comparative effects of low, medium, and high humidity storage
chambers were to be expected, as previous work had already demonstrated
si.mil:tr characteristics in onion seed. But the question rcmains as to
whether alternation of humidities in the storage chambers, and hence
fluctuations of moisture content within the seed itself, is more deleterious
ilian a constant high moisture content. At first it appeared that the
answer to this question was in the negative. Seeds which had two-weekl)',
four-weekly, or eight-weekly ahernations from 35 to 55 per cent rebtive
humidity remained viable longer than those at a constant relative humidity
of 55 per cent, and deteriorated more rapidly than those at a constant
relative humidity of 3S per cent, Seeds kept under conditions of altern:ltion of humidity, then, were intermediate in germination capacity between
those kept at the two humidities concerned. This same response applied
to seeds alternated between 35 :lnd 76 per cent relative humidity and
beLween 55 and 76 per cent relative humidity. However, when these last
two alternations were made eight-weekly, the seeds deteriorated ;IS

.6

MOISTURE EllFECfS

I'apidly as they did when kept constantly at the higher humidity. This
was noteworthy in vicw of the fact that the seeds had been kept for half
of the time at either 35 or 55 per cent relative humidity, both of whieh
permit retention of viability for fairly long periods, and consequendy had
remained for only half as long at the n:ry harmful 76 per cent relative
humidity as the control lot at this humidity.
Further examination of the deterioration effects after certain storage
periods revealed that still other factors were involved. The data shmyed
that during the first nve!ve weeks of storage at nm.. . or four-weekly
alternations of either 35 to 76 or 55 to 76 per cent relative humidity, the
lower humidities tended to prevent rapid loss of viability during the
periods at 76 per cent. However, at the sD.1:ccn-wcek storage period and
thereafter, the lower humidities could no longer counteract the deleterious
effect of the high humidity, and the genmnation-capacity curves morc
nearly approachcd that of the constant high humidity. In other words, at
sixteen weeks of storage and thereafter, deterioration at alternating
humidities was at a more rapid race than would be expected from the
behaviour at each of the humidities. This was especially marked for the
alternation of 35 to 76 per cent.
On the other hand, at an alternation of;;5 to 55 per cent, the periods
at the favourable rclative humidity of 35 per ccnt were sufficient to
decrease the deterioration rate from that expected on the basis of the
behaviour at 55 per cent for up to forty-cight weeks of storage-after
wruch the same cffect as mentioned above was indicated by tests wade
after fifty-two, fifty-six, and si..'Ct}'-four weeks of storage. This delay in che
ipanifestation of the harmful effect produced by alternation of moisture
content was no doubt due to the fact that 55 per cent relative humidity
was much less harmful co germination capacity than was 76 per cent.
The eight-weekly alternations deserve special considcration. The first
viability test of these seeds was made after sL'{tt..'Cn wt..'Cks of storage. As
all seeds in alternating humidities were placeu in the lower humidity first,
this means that seeds at 55 to 76 per cent alternation, for example, had the
first eight weeks at 55 per cent (which permits high retention of viability)
and the second eight weeks at 76 per cent. In spite of the facc that these
seeds had onI)' eight weeks of the unfavourable 76 per cent relative
humidity, after which, if no other factors were involved, they should still
have givcn 67 per cent germination, they gave only 14 per cent germination. A similar effect was obtained after eight-weekly alternation of 35 to
76 pcr cent relative humidity, when the germination obca.ined was 29
per cent.
"'hen the tot:1llength of time at the higher humidity-whcther thac
time was obtained by altemation or by constant humidities~is considered,
the gennination percentages parallel the length of time except ill the case
of the eight-,,eekly alternation which gives a much lower germination
percentage as described abovc. The total period at high humidity, then,

27

SE)

I>R.ESI::R\'ATION A.c'\n LONG\oITY

is of great importmcc in determining the length of life of onion seeds,

but intervening periods of lower and hence more favourable humidities
do not prolong the life-span as much as would be e:xpecre.d from the
behavioural data of seeds kept constandy at these lower humidities.
If the tota.! period at 76 per cenf relative humidity in alternating
humidities were the only facror involved, two-weekly, four-weekly, and
eight-weekly alternations should bring abom the same deterioration when
the total time at 76 per cent is the same. Also, an alternation of 35 to 76
per cent relative humidit) should be as deleterious as one of 55 to 76 per
cent, Such was not found to be the case. Therefore, some other facror
must have been operating, and it appears from the data at hand that this
factor was Oucruation in moisture content.
Dandelion seeds were similar to onion seeds in their response to
moisture fluctuation (Barton, 1943). The length of time at a harmful
relative humidity of 76 per cent was directly related to the deterioration
rate of dandelion seeds, regardless of the storage conditione; before and
after this e.'q)OSlU"C.
ARTlJ1ICIAL DRYING

As the moisture coment of seeds is of prime importance in determining their keeping quality, and as most seeds containharmful amounts
of moisture at harvest-time, it becomes necessary to adjust the moistureto a safe level before storage. Various types of dryers for vegctable and
herbage seeds have been described by North (19..;.8). He places the
maximum air temperarure for safe drying at not more than ~20"F., while
some seeds, such as those ofbeans, onions, and leeks, should not be exposed
to temperatures higher than 90F. during drying. Christidis (1940), on
the other hand, found that cotton seed can withstand the effects of dr:;ng
at 60C. for at least eleven hours. and at 90C. for two to fOUT minutes,
without loss of vitaliry. He recommends 70 to 7SC. as an appIO'priatc
temperature range for drying these seeds. The time required (usually a
few minutes to half an hour) to eliminate the excess moisture in cottonseeds depends on the temperature and speed of the current of hot air used,
as well as on the amount of moisture to be removed. Seeds of crimson
clover (Trifolium inrornalum) can be cured, either by natural drying or by
artificial heat of 43'3C., to a moisture content of 10 per cent ('Vard &
Butt, 1955). Brewer & Butt (1950) dried seeds of the blue lupine (Lupinus
angustifolills L.) at 1 ISO to q.oF. (46 to 60C.) in a forced-draught m'en.
Artificial drying of maize "ith heated air under forced-draught conditions
is used extensively by growers of hybrid seed, according to Kiesselbach
(1939) Under good management this practice can remove the danger of
injury from freezing and also facilimte early harvest, processing, and
storage. Kiesselbach recommended a reduction in moisture content of the
seed to IZ to 13 per cent at a temperature nmgeof 105 to 110F. (40" to

28

MOiSTliRE EFFECTS

.1fC.), but when the initial moisture content is n'-'ar 50 per cent, the
drying temperature should be held to 105F.
Sulphuric acid is a good drying agent, but has the disadvantage that
fumes from the acid may injure the seeds. Also, it is impractical to use.
Glycerine or calcium chloride, in either case mixed "ith water, have also
been used (Nakajima, 1925). In addition to calcium chloride, Kondo
(19260.) used lime, wood ash, and straw ash for mixing with seeds to
control the moisture content.
Seeds may be dried br being placed over or mixed with such drying
agents as calcium chloride, calcium oxide, or silica gel. Some specific
effects of sueh drying methods "ill be discussed in the chapter on vegetable-seed storage. Gennination as affected by storage over various
hygroscopic substances has been described by Nakajima (1927).
For larger seed-lots, and especially for the scedsman who must maintain a high degree of viability in any surplus seed stocks, a dehumidified
storage room is ofren useful in practice. The importance of the temperature in the storage chamber should not be overlooked, and this aspect
will be discussed in the next chapter.
The advantage of pre-harvest drying of rice by chemicals sprayed
from an airplane has been described (Anon., 195311).

IV
OTHER FACTORS AFFJ::l.IING LONGEVITY
TEt'l.fPER...\Th'RE EFFECfS

Jusr as it is impossible to discuss the efl:ect of moisrure contcnt.on the

viability of seeds without considhation of the temperature, so it is impossible to list temperature effects \Yithout consideration of the moisture
COntent. However, the data secured over a period ofsc\'cral years by many
workers in the field show certain trends which :lrc attributable to tcmperahire.
In general, the higher the temperature the more rapid the deterioration
at a given moisture level. Conversely, the lower the temperature the
greater the toler:lllce of high moisture content-apart, of course, from
the mechanical injury of the seed tissues at temperatures below freezing.
Most often, experiment--" set up to show temperature effects have induded
only two or three rcmperarurcs; these were usua1.1,.v ordinary room
temperature, which varied with the locality, and a lower temperatute
approximating to that of a cool cellar (S" to lO"C.).lf the moisture content
of the seed is not too high) i.e. if the humidity in the cold-storage room is
low, maintenance at 5" to woe. will e:\,'tend the life of me seeds far beyond
that under similar humidity conditions at ordinary room temperature. A
high humidity room at Soc., however, will bring about much more rapid
deterioration of seeds than a dry room at 25C. This has been demonstrated repeatedly and will be discussed under methods of storage of the
seeds of individual species.
Perhaps the most significant finding of recent years regarding the effect
of temperature on seed "'lability is the discovety that below-freezing
temperatures are superior to above-free~ing ones for keeping se,,-eral
different kinds of seeds. Only a few instances will be cited here for illustration; others will be discussed under the individual species.
Conifer seed storage may serve as an example. The general literature
on this subject will be reviewed in the chapter 011 tree seeds (Chapter IX).
'Ve are concerned here only with the evidence for the superiority of
belOW-freezing temperatures for storage of these seeds. One of the first
controlled experiments ''laS that of Rarron (r9350)' who placed conifer
seeds under different conditions at three temperatures: (1) room, (2) Soc.,
and (3) in a refrigeration room with an average temperature of -IS~C.
for the first two or three rears of the cxperimenl<;, afrer which they were

30

OTHER FACTORS AFFEO'ING LONGEVITY

transferred to another refrigeration room with an average temperature

of ~ Soc. The seeds used were of loblolly pine (Pillus laeda L.), slash pine
(Pillus cnribnen Morelet), shortleaf pine (Pillus ec1/iwlfa Mill.), longleaf
pine (PiIllH paluslris Mill.), red pine (Pinus Tesillom Ait.), Pinus P01lderosa Douglas, Picea exce/sa Link, and Picea glauca (Moench) Voss (P.
c(madel/sis of authors). The conclusion at the time of the first report of
the results of these tests, when the seeds had been in storage up to seven
ycars (Barton, 1935a), was that scaled storage at either of the low temperatures (SoC. or -So to -15C.) was effective for the maintenance of
viability. However, the data showed a significant superiority of the lower
of the two temperatures at that time ('Vakeley, 1945). Further tests
(Barton, 1953(1.) showed that some of these seeds relll3.ined viable for up
to sixteen years, when the supply of seeds became exhausted. Isaac (1934)
stored noble fir seed successfully for five years at ISoF. (-TOOC).

TABLE VII
GER.\lli.... ATION M'D MOISTIJRE COi'<TI':NT

CONIFER SF.F.DS AFTER

STOR.,I,GE AT SUll-FREF21NG TF_\fPERATURES

Species and pef cent

germination at time
of stomge

Approximate
storage
temp. ("c.)

-,

Ponderosa pine

-H

95

"

-,

Douglas fir
88

-H

-"

-,

Sitka spruce

,6

-H

-,8

-,

Wesrern red cedar

60

-H

-,8
Western hemlock

-4

-,8
-

p" cent germination after storage fo,

I, 2,

87
8,
8,

9'

9'

79
75

7'
7'

'"

44
75

8,

88

8,

"6,62

66
6,
74

5
7
'9

Tlldic:ltes thai no le<;I

9'
79
77

8,
8,
74

33
6,

"6,

3, 4, and 5 yr.
3

38
55

60

78

,
"47

,
0

8
43

",

",

76

'4
6,

70

0
0

0
0

"

'3

27
3'

Wll.~

9
'-l

rrulde.

With this knowledge, and because of a need for determining the best
possible storage condition for valuable, short-livcd conifcr seeds, a test
was started in which three different sub-freezing temperatures were used
for storage of seeds of Ponderosa pine (Pillus ponderosa Dougl.), Douglas

3'

SEED PRfSERV,UlON AND LONGEVITY

"fir (Pseudotmga ta.l'ijolia Britt.), Sitka spruce (Piceo. sitclJemis Carr.),

Western red cedat (Tlwja pliClllfl Donn.), and Western hemlock (Tsllga.
he/crop/gila Sarg.) (Barton, 1954)' The :J.\'ailable seeds of each species
were divided into three lots which were stored in canvas bags at temperatures of approximately - 4", - II", and - I BeC. For the first two temperatures, cold rooms were used with average temperatures as indicated
but ",':ith a fluctuation of 5"C. Afood freezer furnished the more constant temperature of -18"C.
"Or-------~-------~-------,

'0

~
~

60

W
0

40

Storage
Temperature

w
~

Ge
-I f'c
-I aGe

'0

-4

0
0

Douglas Sitko
Fir
Spruce
00

,
YEARS OF STORAGE

FIG. 6.----GraJ?h indicating pero:ntagc gcnnin:uion of seeds of Douglas fIr and Sitka spruce
after storage at sub-freezing temperaLures.

Viability tests were made at the beginning of the experiment and then
annually for live years in succession. Results are shown in Table 'VII.
There was great variation in the ,,;abiliry of the different species at the
beginning of the experiment. The seeds of Ponderosa pine and Douglas
fir were of excellent quality, giving 95 and 88 per cent germination,
respectivel)', at the time storage was begun. The seeds of Sitka spruce and
Western red eedar were less vigorous, with initial germination perccntages of 56 and 60, while only 12 per cent of the Western hemlock
seeds germinated.
All of the seeds, except those of Ponderosa pine, 'kept' better a[
-ISDe. than at -IIC., and deterioration was most rapid at -4"C.
(Table VII). It was anticipated that some such effect might be obtained,
but the rapidity of the response was striking. Definite differences in
deterioration tates were evident after only two years of stor.tge. In
cxperiments with other seeds (Barton, T9531J) at least five rears of storage
32

OTHER FACroRS AFFE(.IING LONGEVITY

were required to show differences in response between 5 and _+0C.

Tbc germination capacity of Douglas fir and Sitka spruce secds aftcr
storage at the three temperatures is shuwn graphically in Fig. 6. It appears
that the lower the sub-freezing temperaturc (to -18"C.), the better the
survival of Douglas fir and Sitka spruce seeds in canvas bags.
Other tests (Barton, I954Jl) have shown that reduction in moisture
content extends the life of Doughs fir and Western hemlock seeds e....en
at the very favourable temperature of -IBC. Allen (1957) found oF.
better than 32F. for storage of conifer seeds in only a few cases out of
twenty-seven lots representing eleven species. He stored thc seeds in
sealed vials for periods of from five to seven )'t..':lrs.
Although these conifer seeds, together with maize and with vegetable
and flower seeds (see chapters on these various seeds), have been benefited
by storage below freezing, it should not be assumed that all seeds are so
al1ccted. Weibull (1955) points out that, while cold storage at -20C.
is good for maintaining the viability of many kinds of vegetable seed, it
cannot be used for parsley (Petrusclin1l11l sativum) seed, which reacts
adversely to cold storage and loses germination ability rapidly at -20C.
Also, the seeds of four types of flower, namely A'lli"lJirlUtII majllS, Aspamgus plt/mosus, Peflm; Irybrida, and Viola fricolor var. maxima, are
actually killed at -20<>C. As a result of his experiments Weibull (1955,
p. 667) concluded that 'a test should therefore be made before the largescale application of cold storage to a certain kind of seed'. These results
notwithstanding, the discovery of the successful storage of many types of
seeds at sub--freezing temperatures eliminates the necessitr for scaled
storage and maintains a high quality in the sl-eds which a~e then able
to resist further unfavourable stOlage conditions (see Chapter XI, on
packeting, etc.).
It will be noted in Chapter VII that grains, especially maize, are now
being stored at sub-freezing temperatures.
GAS EFFECfS

Gaseous exchange is important in the merabolism of the seed and

hence has a relation to its life-span. Undoubtedly respiration is affected, but
the factors involved in seed mortality appear to be much mOle complex than
those in ordinary aerobic respiratioIi. In any case, the gaseous exchange
is direcdy related to the moisture content of the seed and to the temperature at which it is stored. Because moisture and temperature conditions
seem to be mOle directly concerned in deternUning the life-span of seeds,
less experimental work has been done on the effects of different gases.
A detailed experiment on cotton~seed storage in various gases under
controlled temperature and moisture conditions has been reported by
Simpson (1953). Initially, moisture content of the seeds llsed in this
experiment was 10 per cent and the germination was 97 per cent. Moisture
c
33

SEED PRF.SERVATION A,\I-n LONGEVITY

was adjusted to two levels, 7 per cent by drying in the sun and 13 per
cent by adding water. The seeds were then placed in pint milk-bott1es
fitted with tight cork stoppers. The corks were pn.'Ssed down below the
rim of the bottle, 1e-<1ving room for a paraffin seal. The gases used were
air, o~ygen, carbon dioxide, and nitrogen. The last three gases were
introduced into the bottles by evacuating the latter and then flushing them
with the desired gas t\vice before filling, adjusting the pressure to that
of the atmosphere, and sealing with paraffin. Each bottle contained
approximately 100 gm. of seed. The total volume of gas in each bottle
was about 355 ml.
Examination of the samples was made every six. months for three-anda-half years. After that time, lots with 7 per cent moisture and at 7o"F.
were held in reserve and tested aftt.'! eight and ten years of storage. Gas
pressure and \\eight of free Cilrbon dioxide within the boule, and moisture
content and germination percentages of the seeels, were obtained from
duplicate samples on each sampling date. Chemical analyses were also
made on seeds stored for two-and-a-half, three-and-a-half, and ten years.
Variation in moisture content during the storage period did not
exceed that which would be expected from sampling errOr. Seeds containing 7 per cent of moisture were still fully viable after storage for ten
years at 70"F., and there was no deterioratio.n of seeds with this moisture
content at 90"11. after three-aud-a-half years, when the supply was exhausted. At the higher moisture content of 13 per cent, viability was
retained for si.x months at 70F., but deterioration was rapid after I.hat
time,and only a few seeds were capable ofgerrnin:ation afterone-and-a-half
years. At 90F., seeds with 13 per cent moisture failed to survive for as
long as six months.
There was a decrease in gas pressure in the bottles with seeds sealed
in air, oxygen, or carbon dioxide-most logically explained, according to
the author (Simpson, 1953), by assuming that carbon dioxide was
absorbed by the seed. Containers with atlllospheres ofnitrogen gas showed
little change in pressure with storage condition or time. The carbon
dioxide recovered from the containers after intervals of storage indicated
rapid production of carbon dioxide by the seeds in an atmosphere of
oxygen, moderate production in air, and low production in nitrogen.
Considerably less carbon dioxide was recovered after storage than was
placed in the containers at the beginning of the test, i.e. the seeds must
have absorbed carbon dioxide. The author concluded that, within the
limits of this experiment, the initial atmosphere in the storage chamber
had no effect on the longevity of the cotton-seed, and that free o:-.."ygen
was not essential to the continuous life-processes of the seed. Therefore,
moisture and temperature of storage are more inlportant than atmosphere
in keeping cotton-seed for ten years.
Sayre (1940) sealed maize grain for five years in air, oxygen, carbon
dioxide, and nitrogen. There was no deterioration in samples stored at

34

--

OTHF.R FACTORS AFFEcrll'\G LOl'\GVIn'

3C. and -z5"C. At 30C., however, deterioration ,,-as c,;dent in all

samples after storagc for three years, but loss of germitl:J.tion capacity \\ as
more pronounced in O:'l:ygen and nitrogen atmospheres than in air or
carbon dioxide. As a result of these tests, the author questioned the theory
that free oXygen is necessary for continued ,,;rality of stored seed, and
stated that respiration may speed deterioration and consequently any
condition which will decrease respiration will extend the life of the seed.
The reason for the harmful effect of thc nitrogen atmosphere on stored
maize breain is not clear.
Busse (1935) stored poplar seeds in air and in a partial vacuum and
found that the life-span was increased by reducing the air pressure. He
believed this to be due to the injurious action of the oxygen in the air.
Guillaumin (1928) demonstrated by an experiment lasting twelve years
that a vacuum was a means of presen;ng the viability of grains. An
atmosphere devoid of oxygen was as good as a vacuum; hence the harmful
effect of oxygen itself was shown. Sampietro (1931) found that nitrogen
extended the life of rice seeds in stomge, and that this gas was better than
carbon dioxide, air, or a vacuum. Air and carbon dioxide had the same
effect on rice in storage, according to Kondo & Okamura (r93-1).
Seeds of the American elm stored in an atmosphere of o:\l-gen lost
their viability more quickly than others sealed in air. When moisrore or
temperature conditions were unfavourable, a partial Yacuum extcnded the
life of these seeds (Barton, I93C}1l), Some benefits of a lessened oxygen
supply Iruly have been experienced by conifer seeds under certain storage
conditions (Barton, 1935a), but aster and Verbmaseeds were not improved
by a partial vacuum undcr the conditions tried by Banon (19530.).
The fact that an atmosphere of pure oxygen increases the respiratory
ratc of seeds, leads one to expect that this gas might shorten the life of
some types of seeds under cerrain conditions-especially those of unfavourable moisture and temperature~and perhaps more significantly
for seeds of low germin:ltion capacity than for others. Furthcr evidence
of the harmful effect of oX)-gen bas been demonstrated for barley seeds
stored after X-radiation (Adams d al., 1955)' It was found that o:\1'gen
enhanced the amount of radiation damage as measured by frequency of
chromosome bridges, rate of germitl:J.tion, and emergence of seedlings,
while nitrogen retarded these effeclS.
It seems certain that most seeds can carry on the minimum metabolic
activity necessary for retention of viability without an outside supply of
oxygen. and hence would not be benefited by such a supply. Ohga (19260)
found, for example, that internal gas pumped out of very old but viable
Indian lotus fruirs was composed of an average of 18'33 per cent oxygen.,
0'14 per cent carbon dioxide, and 80'93 per cent nitrogen-as compared
to 1888 per cent O:'l.ygen, 0'81 per cent carbon dioxide, and 80'31 per
cent nitrogen in fresh fruits. The permeability of the seed-<:oat to gases
is at a minimum in dry seeds and increases with moisture (Becquerel,

35

SUD PRI.SERVATIO:-: A.''D 1.o.,\,GEVITY

1904). This may well be

factor in the favourable effect of drying for

many types of seeds. The whole matter of permeability is an importanl
one, but is outside the St:ope of this book.
3.

LIGHT

EFt'"ECI'S

Although there are numerous publications concerning the effect of

light on the germination of seeds, there is very little available information
on the effect of light on seeds in storage. No effect on germination of
ex-posure of eight different seed-types to the sun for forry-four days was
noted by Tammes (1900). \\'hen the moisture content of seeds of Gil/rhol/a
JedgeritJlltJ was as high as 9'4 per cent, there was, however, some e,,-idence
of a harmful effect of exposurc to light during storage in the laboratory
(Barton, 1947).
I
C. Jensen ([941) treated seeds of cauliflowcr by exposing them to a
Qp.artz-lamp and a Sollux-lamp together, n-irh full strength for one hour
at a distance of 1 ffi. from thc lamps. The seeds were stirred frequently
during the exposure. Following this treatment the seeds were stored in
glass containers plUb'ged with an ordinary cork or a paraffined cork.
Light-treated seeds maintained a higher germination percentage ovcr a
period of eight years thm untreated seeds. Light treatment not only
e:\1:ended the life-span, but also increased the germination capacity and
vigour of fresh seeds. No moisture determinations were made before or
after treatment, and it is possible that the light-treatment procedure may
have reduced Lhe moisture content to a more favourable level. In any case,
the whole subject of the effect of light during storage is a field which
should be exploreu morc thoroughly.
L"-'HERtrA."O:

]. W. Jones (11)26) noted that the seeds of some rice varieties appear
to deteriorate with age more rapidly than others. Three-yca.r--old seeds
of five varieties ranged in germination from 85 to 99 pcr cent, while U10SC
of three other varieties of similar age gcnninated only from 19 to T~ per
cent.
Maize seeds homozygous for luteus: and lureus.a genes lose vi:lbility
more quickly than seeds homozygous for any of the remaining sixluteus
genes (Weiss & Wentz, 193j). Fresh seeds wlth luteus 2 and lutcus", genes
germinated as rapid!)' as nonnal seeds, bur such luteus seeds when a year
old germinated much more slowly and the plants produced from them
also grew more slowl)' than normal plants. The authors suggest lhat it is
possible that these genes do nOt cause decrease in gcrminability, hut that
each of these two genes is closely linl:cd with another gene which causes
loss of viabilit)'.
Lindstrom (1942) also found that embryo longevity of maize is heritable. lie made germirutimL tests with hundreds of inbred lines and F ,
crosses of fiyc- to n,"d,"e-~'ea.r--oldseeds stored at room temperature. Tht:y

36

on IER

l'ACfORS

AFFECrl..t~G

LONGEVITY

showed a variation in germination of from 0 to 90 per cent. Further tests

were made under controlled temperature and humidity storage conditions.
It wa.,> found that hybrid seed of long-lived by shan-lived parental
inbreds exhibited the long-lived condition, oftcn with significant differences in reciproeal crosses.
Although other examples of detailed experiments to determine the
inheritance of longevity arc lacking, it is obviolls that the general make-up
of the particular seed, especially the permeability of the coat, is a major
factor in determining its life-span.

SEED-COAT EFFEcr.s
The importance of the seed-coat in the life of the seed was recognized
early and has already been discussed (Chapter I). Rces (19II) gave
evidence that the impermeable part of the coat was a waxy curicle in many
species, and the thicker the cuticle, the longer was the time in sulphuric
acid required to produce swelling when treated seeds were placed in
water. Wahlen (1929) found that longevity in dover seeds depended upon
the impermeability of the coat.
Mechanical injury to seeds by threshing usually contributes to immediate reduction in germination capacity and to an accelerated loss of
viability in storage. Certainly the embryos of such seeds afC morc vulnerable to attack by micro-organisms. Scarification of alfalf.'l seeds (Battle,
1948), mechanical dehulling of bromegrass seeds (Ahlgren et al., 1950),
and dclinting of cotton-seeds with sulphuric acid (Hamid, 1938), arc all
examp!c... of treatments which reduce the life-span of the seeds. Leaving
the chaff on stored seeds of several species and stocks of wheat and two
varieties of barley, appears [Q be the bestfor storing; but such sl.'Cds should
be shelled before testing or planting, as removal of thc chaff results in a
considerable inctease in the percentage of germination (L. Smith, 1948).
On the other hand, Cutler (1940) found no indieation of loss of viability
attributable to rubbing or clipping oat grains.
Although thc seeds of the longest known life-span are those which
have remained impermeable during the storage period, it is sometimes
important for planting purposes [Q have the seeds become permeable
during storage. Conversely, seeds which are permeable when harvested
may become impermeable during storage under certain conditions of
temperature and humidity. Whilfi such 'hard' seeds are oftcn still alive
when tested, there are some in which a high percentage are not viablee.g. Vicj{l villQsa (V. K. Toole, 1939). There is a considerable amount of
literature on the development of hard seeds in storage, bur the subject
will not be discussed further in this book.

MATURITY EFFEcrs
Studies on the effect of maturity on the germination of seeds have been
made principally to determine the proper date for harvesting of seed for

37

SUD PRESERV.'\110N Al\.'D LONGY....'IT

sowing. The germination tests have been related to the performance of the
seeds immediately after they had been han-csred, rather than to their
resistance to subsequent storage.
Seeds of ion-ay spruce and Scots pine ripen prior to the lignification
of the cones, and hence cones lurYest~d early ,,-il\ dry Out morc rapidly
than those from a later harvest. Rapid dry;ng accounts for the Im\er
germination after a year of storage of seed from cones harvested early,
according to G. Vincent & Freudl (1931).
To stuay the effect of maturity on the '-iability and longevity of the
seeds of western range and pasture grasscs, McAlister (1943) collected
seeds of AgropyrolJ cristlltmn, A. smithii, A. tra&hycaulum, Brot1lus inermis,
B. marginatus, B. /J()1yo1l1/ms, Elpnw glaru:us, and SliptJ viridula, in the
pre-milk, milk, dough, and mature stages of de.elopmcm. Greenhouse
tests of the germination capacity of these seeds were made after storage
for +. 9, 15. 22, 40. 51, and 58 months. Pre-milk and milk seeds were
inferior in germination and longevity to seeds harvestc.d in cither dough.
or Jll3.ture stages. Exceptions to this were the pre-milk and milk stages of
seeds of Bromus margina/us and B. pO!;'anthus, which gave as high:J. perI;entage of germination as mature seeds during the whole period of storage.
Dough-stage seed had similar viabili~' and longe...-ity to the mature seeds
in all species, when tested in the greenhouse. Tn field plantings, however,
tbe immature seeds were gener:lll~' much inferior, as far as seedling
emergence was concerned, to those harvested at maturity. The only
immature seeds which gave as good a stand as the mature seeds during:
the three years foUo1,\-ing collection were those of the dough stage of
Bromus marginstus. No differences in sunrival or size of plants from matur.e
and immature seeds were evident bv the end of the growing-season in
the field.
Rice seed for storage and sowing should be saved from early and
medium plantings as sueh grain has a lower moisture content at harvest
time than has grain from late plantings (McNeal, 1950). It has been
suggested by Riddell & Gries (195q) that the variations in growth of
spring wheats from seed of different ages arc related to the temper-ature
during maturation rather than to me age of the seed or to conditions
during storage.
DOR~UNCY EFFECfS

A Imowledge of the presence of a dormant period is import:lllt in

developing methods for testing seed viability. Dormancy as expressed by
special requirements for germination makes possible the cxtended life
of seeds buried in soil (see Chapter II). Dormancy imposed by an impermeable seed-coat is a significant factor in seed longevity, as we have
alread)' seen. The oldest known viable seeds arc those with hard seedcoots which resist water absorption. Because of this character, seed-eoats
often cause delay in germination (Crocker, 1906; and others).

38

OTHER FACroRS AFFECTING I.ONGEVITY

Embryo dormancy is also important in germination and hence in tcsts

for viability, but is not directly relatcd to longevity_
FRUIT OR

PeL;) EFFECI'S

Many studies havc indicated the inhibiting cffect of fruit or pulp on

germination, but few authors ha\-c rcported a subsequcnt effect on
longevity.
Pulp disintegration of peaches in a pile of twenty bushels of fruit
teduced the germination of the seeds because of the high temperature
caused by rotting, and also because of some unknown property of the
fermenting juice (Haut & Gardner, 1935). Germination tests of seeds
removed from mature and immature fruits of butternut squash have been
made after 211 days of storage (Holmes, J953). All the seeds germinated

well.

39

v
onn:R FACTORS AFFECTING LONGEVITY (coord.)
MICROFLOR.>\

THE role of bacteria and fungi associated \\il.h seeds in storage in bringing

about their deterioration has been a subject of interest for many years.
This interest has led to much detailed work on the subject. Two excellent
reviews (Semeniuk & Gilman, 1944; Christensen, 1957), and a chapter
on microflora written by Semeniuk in a recent book on Storage ofCercaJ
Grains and t!lt'ir Pruducts (Anderson & Alcock, 19S.i.), should be consulted
by the reader for the present status of the problem, and for a complete
review of the literature on the subject.

Workers at the University of Minnesota have published a series of

twenty-five anicles on grain storage studies from 19+5 to 1957- The titles
of some of these publications indicate special interest in- the microflora of
stored seeds, espcially soya beans, wheat, and cotton-seed O'olilner &
Geddes, 1946, 1946a; .Milner (/ 01.,19_1-7. 1947a; Christensen et al., 1949;
Hummel et Ill., 1954; Chrjstensen, 1955, 1955; Sor;cr-Domenigg et al.~
1955. 1955 a ; Papavizas & Christensen, 1957; Tuite & Christensen, 1957),
Some of the more recent of these studies will now be discussed.
WHEAT

It is to be expected that microflora will grow better on seeds with high

moisture contents, resulting eventually in e..xcess heating and mould growth
when seeds are stored under conditions of high humidity and temperatures favourable for fungus growth. Christensen (1955) inycstigated the
invasion by moulds of wheat b'Tain stored in the 1:lboratory for sixteen
months and containing 13'5 to 15.0 per cent of moisture. He found that
Aspergillus resfric!us invaded grains under these conditions and that the
invasion was associated with decreasing germination and increasing
development of a brown colour of the germ which is characteristic of
'sick' wheat. The main invasion by the funglL'i appeared to be iu the region
of the germ. These results indiC<!.te that a moisture content of L3-S pcr
cent. which has been commonly accepted as a safe one for grain storage, is
too high, as it does perm.it i>10W invasion by this fungus.
In further studies, the effect of mould growth (during temporary
exposure of wheat grain to conditions of high moisture) upon the development of germ damage during subsequen.t stof:J.ge was determined (SorgcrDomenigg et I., 1955). Seeds of 'hard' red spring wheat which had been

OTHER FACWRS AFFECrJNG LONGEVrn

inoculated with an appropriate volume of water suspension of mould

spores to give samples containing 15, 18, and 21 per cent moisture. After
five, ten or fifteen days incubation, the wheat samples were air-dried to
approximately 13 or 14 per cent moisture content on trays at room
temperature, where thcywere then stored. High levels of mould infestation
increased the development of germ damage, and this was accompanied by
loss in germination capacity and by increased 'fat acidity' during the
temporary storage at high moisture levels. During ~;llbsequent storage at
13 and 14 per cent moisture, the mould population of the majority of the
samples decreased, but the germ damage and fat acidity continued to
increase, especially in the samples whieh had been heavily infcsted .,.,'ith
mould. Not only did inoculated samples temporarily exposed to 2I pcr
cent moistute show Joss of viability, bur seeds so treated gave flour of high
ash content, poor colour, poor baking strength, and general loss in baking
quality. Negative correlations were found between free fatty acids and loaf
volume, germ damage and loaf volume, and the logarithm of viability and
free fatty acids. It was pointed out that los,.<;cs in viability preceded the
discoloration of the germ and were indicative of incipient damage and poor
storage properties. The authors srate (Sorger-Domenigg et ai., I955, p.
282): "While not excluding the possibility that sick wheat may develop in
the absence of mold growth, the present investigations provide strong
evidence that molds are largely responsible for the losses in viability which
precede the discoloration of the genn and the increases in fat acidity.'
However, it would appear that the sharp decrease in the mould count in
wheat stored at 14 per cent or less moisture after exposure to high
moisture, even though other tests such as viability, germ damage,
and baking quality indicated continued decline of seed quality,
might argue against the mould invasion as the sole cause of deterioration.
Papavizas & Christensen (T957) made a study of various fungi
associated with wheat seeds in storage, to determine the effect of each
fungus on the development of 'sick' wheat. They stored grains after
sterilizing with sodium hypochlorite (I per cent for one minute) and
drying to a moisture content of II to 12 per cent. They point out that
storage fungi do not innde seeds to any appreciable c),."tent before harvest,
and that wheat seeds sterilized in this war were almost completely free
from fungi. Some of the seeds were, then inoculated with dry spores of
Aspergiiiu5 candidus, A. repellS, or A. restriclU5, and stored for up to seven
months in desiccators at 25C. over humidifying solutions giving moisture
contents of from 147 to approximately 20 per Cent. Non-inoculated seeds
declined very little in geIOunation capacity and produced very few
damaged gcrms ('sick' wheat), while those inoculated "ith Aspergiiiu5
deteriorated much more rapidly. This is illustrated in Fig. 7 taken from
the original article (papavizas & Christensen, 1957). Also, of the three
species of Aspergiiills indicated in the figure,.E1. repem was least harmful to
the stored seeds. Invasion of the wheat seed by fungi was followed by

4'

SEED PRESJ::RVATION .<\I\'D LONGl':VITY

the development of germ d:J.mab'"e or 'sick' wheat as pictured in Plate

(from Papavizas & Christensen, 1957).

10'~~~-:=;:_==:==--'----l
r
~.~'-'-------.""","Noninoc
60

.~

-.
'-.-.

----e"'-A.condi

a--.....--'

---.fc-A.condi
.~resi'ic

~.

40

----.~repen

-----.f--A,re stric

--'

o 8
o

en

60

Cl

::E 40

cr
w

'" 2
~

. ' - - - - '<-A~epen

/'

.- . /
5

WEEKS

II

STORAGE -

~oninoc

16
80%

RH

FIG. i.-Graphs showing germination :md geml d.:I.m~ge of grains of Willet wheat noninoculated and inoculated with Asp~ill"s car:didm, .<1.. uprns, and A. ustrutm, and stored
at 25C. under 80 per cent rdati,-e humidily (moil'ture coment t6o to 16'4- per cent) (from
Pap:l\'izas & Christensen, 195i; courtesy of Departmen[ of Plant Pathology and Bot:mY.
University of:Minncsot:I. St. Paul, Minn~ U.S.A.).

Fusarium mmliliforme appeared to bring about more rapid deterioration

of stored seeds of barley than did species of Aspergillus and Penicillium
(Armolik et aI., 1956). These studies were carried further by treating
barley seeds with twenty-day-old culture filtrates of several fungi. A.l.1 of
the filtrates, except those from Pellicill;um c1lrysogenu1II, reduced early
seedling growth (see Plate 3). Fusar;ulII moniliforme, which caused the
worst damage to seeds in storage, also had the most deleterious effect in
filtrate form.

--

an-U::R FAcrORS Atl'-'ECfING WKGEVITY

Although Armolik et 01. (1956) concluded that the fungi were important
factors in the deterioration of moist barley seeds, they also noted deterioration of the sterilized seeds, thus indicating that fungi arc not the only
factors concerned. Further tests on the humidity requirements for
germination of the spores of fungi associated with grain, showed that the)'
would develop on substrates at moisture contents of between 10 and 20
percent, depending upon the fungus (Armolik & Dickson, 1956). Members
of the Aspergillus glaucus group germinated under lower humidity
conditions than did the others, and they were usually the ftrst fungi to
invade cereal grains in storage. Fusarium mOl1i1iforme, on the other hand,
required 204 per cent of moisture for germination, which was the
highest of any fungus stUdied. The moisture content of freshly 'combined'
baric)' grain was found to be high enough to support b'Towth of all of the
fungi studied-which indicates the need for artificial drying, as deterioration is very rapid during the first ten days after harvesting.
To determine the time of invasion of the wheat seeds by forms of
Aspergillus responsible for deterioration in storage, Tuite & Christensen
(I957) collected grains from different varieties and classes of wheat from
several different states over a period of three harvest seasons, and determined the presence of storage fungi by culturing the seeds on malt-salt
agar. Only a small percentage of the seeds were found to be infected.
However, StOrage fungi invaded thc secrls readily when lhey were stored
in a humid chamber, the threshed seeds being more susceptible than seeds
left in the heads. The percentage of wheat seeds ~ielding storage fungi
increased between harvest and the arrival of the grain at the storage site.
Furthcr evidence of the causal relationship of mould to seed deterioration was furnished by Christensen (1955a) in his study of the incidence of
germ damage in commercial wheat grain. Christensen's abstract of his
fin~isasfollo~~:

Gcnn:iJu.rion of the seed, and number and kinds of molds present, were determined in 'sick' and sound seeds picked from 26 commercial samples oontaining
from 5-55 per cent 'sid:' wheat, and in sound wheal from bull:s in which no
detcri0r2[lon had occurred. The germination of 'sick' seed alv.-ays was :terti;
molds wcre microscopically \":isiblc on the gl':nI1S of 49 per cent of the seeds; tbey
had an a\'Cf2ge mold count of402.,ooojg.; and 94 pttceIlt ofLhe surface-disinfceted
seeds yielded slOrage molds. The 'sound' seeds picked from the lots in which
deterioration had occurred had an average germination of 43 per cent and an
average mold count of 32,ooojg., and 84 per cent of the surface-disinfected seeds
yielded storage molds, The really sound seeds from bulks in which no dctcrior:ltion had occurred had an a\'crage genninarion of 9r per cent, an a\'crage mold
count of less than I,OOO/g., and 27 per cenl of the surfacc-disinfected seeds yielded
storage molds. The lll3ior fungi present in the 'sid:' seeds were Asprrgil/us
resrriwn, A. r~jJnIs, A. eandidus, and A. 1=. Judged b)" \-arious microscopic
and culturallechnics, all samples of 'sick' ""beat had been \"try beavily in\llded
by stonge molds; all of the evidence indialcd tb:I.l im'asion of lbe gams of the
seeds by these molds had preceded decrClSC in germination and increase in 'sick'
wheal. In commcrcia.l storage, it seems \"ay probable tJut invasion of the germs
or the seed by common species of As~rgillus is a common ausc of 'sick' wheal.

43

SEED PRESERVATION AND LONGEVl1Y

Some measurements have been made of the chemical changes raking

place in mould-infected as compared with uninfcctcd wheat grain.
Increased respiratory rate, increased fat acidit)', and- decreased nonreducing sugars, were associated with thc presence of mould (Hummel et
(/1.,1954). Loss in viability accompanied these changes, though at moisture
levels of 18 per cent or above none of the seeds survived.
Cole & l'vlilner (1953) presented evidence that the discoloration of the
germs of 'sick' whcat grain is due to a browning reaction of the Ivlailhrd
type. They found a positive correlation of 0'748 bemeen the fluorescence
of aqueous extracts of the sample and an e,'aluation of germ damage by
federal inspcr:tors. They also found a significant negative correhtion of
0'775 between fluorescence and germin.ation..capacity o[I.he samples. They
suggested that the fluorescence tes!.'ctmld be standardized as an. index of
viability.
Sorger-Domenigg et 01. (l955) did not find such high correlation
coefficients between fluorescence value and germ damage or germi.nation
capacity. The correbtion between germ damage and formazan value
(see Chapter XIII), though somewhat higher, still did not justify its use to
predict the value of wheat seed. However, there was a high correlation
(~O92) between viability and fat acidity. B. Thomas (1941) pointed out
that destruction by bacteria can ocwr in cereals with moisture contents
of over 17 per cent.

-:

OTHER SEEDS

Although the presence of many fungi, among which species of

Pellicillium, Aspergillus, and Rlli:.:,opus are predominant, has been demonstrated on stored maize grains, it is not thought to be of sis'Tlificance in the
storage problcm-ehicfly because of the high moisture requirements of
these fungi. Maize is usually dried to a moisture content below that required
for the growth of the fungi (approximately 12 per cent) before it is stored
umler conditions which prevent atmospheric condensation (Semeniuk &
Gilman, 1944; Semeniuk et al., 1947).
Experiments of Venkatram (1950) showed that fungi were not the
primary cause of cotton-seed deterioration, which was due rather to the
high moisture content of the seeds and to unfavourable temperatures
during storage.
Kurata et of. (1957) fuund that micro-organisms, including fllll~,
bacteria, yeasts, and actinomycetes, were not normally present in milled
rice producet.l commercially in the southern area of the United States in
1954
Whether fungi playa major role in the deterioration of seeds in
storage still remains a question in the minds of many seed experts. The
fungi grow only under high humidity conditions which favour deterioration of seeds not obviously invaded by fungi. Furthcrmore, although low
tcmperature and low moisture content prolong the life of mmy kinds of

44

1:3

OTHER FACTORS AFFECTIKG LO;-;GEVTn'

seeds, the embryos finally do die under these conditions-without mould

b'Towth, ,\ ruch is inhibited by dte storage condition.... Also, it has been
shown alxwe tbat sterile wheat seeds deterior.ne under conditions in
which mould invasion accelerates the deterioration. It becomes a question
of wbether the mould docs not attack the deteriorating seeds. This theory
is strengthened by the fact that most of the fungi found on stored seeds
arc saprophytes. It is an invariable rule that seeds of high quality can be
germinated on a moist substrate without infection of the culture by
bacteria and moulds. Germination is rapid and the seeds and seedlings
remain <lpparcntly clean. However, seeds of the same lot which have been
in sealed containers under conditions that cause deterioration but do not
permit fungus growth, will rapidly become mouldy when tbey arc placed
on the germination substrate. Undoubtedly, both seeds harboured the
same microflora at the beginning of the storage period. Whether the
sound seeds produce a chemiCl! deterrent to mould growth, or whether
sugars or other fungus nutrients leach out from deteriorating seeds, is a
matter for future tests to determine.
In cold tests used to evaluate some of the factors contributing to the
longe,,-ity of maize seeds, it has been noted by Goodsell t1 al. (1955) that
there is an increasing susceptibility to soil-borne pathogens W1th an
increase in age of the seeds. They nate that this phenomenon is not well
understood but may perhaps be related to penneability changes.
A knowledge of biochemical changes which take place in deteriorating
seeds under dry conditions at low temperatures over long periods of time,
as compared to those taking place under moist conditions in a short rime,
would aid greatly in the interpretation of the role of microflora in the (oss
of Y-lability by seeds.
LIFE-SPA>" OF Ft.iKGI

The life-span of the fungi associated with seeds in storage is a matter of

some interest. It is not "'-lthin the scope of this book to discuss longevity of
fungi in detail, but a few examples will be given. The longel-l)' of two
fungi occurring on wheat seeds was compared by Russell (l956), who found
that whereas A/umaria tenui! disappeared after seven p.. a rs of storage,
S per cent of the He/l1Iinthosporium sa,tmum was still alive after seventeen
years. The seeds themselves retained their full viability for ahom nine
years under the conditions tried by Russell, thus indicating that prolonged
storage of the seeds could be used to eliminate the Altermma but not the

He/minthospnriulIl.
Machacek & Wallace (1952) stored wheat, oat, and barley seeds,
infected with fungi, in paper hags in galvanized metal hoxes at room
temperature. At intervals up to ten rears later, samples of the seeds were
surface-SIerilized and tested for the presence of the fungi. Alter11l1ria
tmuis (sensu Wilshire) in all three types of seeds. Helminthosporium
SlJlr';U111 P.K. & B. in wheat and barley grains, and Septorill nodorum Berk.

4;

SEED PRESERV.4.TION .o\I'\'D LONGEV.1'l'Y

in wheat g~ilJ.S, all detcriomtcd rapidly. after' one year of storage.

HelmintllOs'1,(iruiln. ll'vmue Eidam i'n oats add H.-:ures Sacco in bark}.",
deteriorated more slowly, but a shall percentage of each of the fungi was
recovered after seven to ten years of storage. The pathogens remaining
alive at the end of ten years when the seeds were still viable were considered to be ofsmall consequence as far as crop production was concerned.
E. Brown & Robert (1943) found that Alternaria sp. on oats, wheat.)
and barley seeds were killed when stored for from six to fifteen months at
105 of., although the seeds would still germinate. This treatment should
be used with caution, however, as seeds so treated may not be vigorous
enough to produce a good crop, and prolonged storage at such a. high
temperature would certainly result in their death.
In rice seeds containing 18 to 26 per cent of moisture which had been
sealed for sevcn months, SOme moulds survived but did not increa.<;e,
aerobic bacteria remained the same or decreased in number, while
facultative anaerobicorganislIlS and yeasts increased in number (Teunis.'mn,
1954) There was a loss in viability of the rice seeds during this period.
Ageing of maize seeds reduced the prevalence of viable fungi in infected
seed grain (Dungan & Koehler, 1944). During the first year of storage,
Nigrospora oryzae infection was gready reduced, Diplodia zeac was
reduced about 50 per cem, and Gibberella zeae was almost completely
eliminated. Although Cephalosporium acremollium and Fusarium monilifonne
infections were reduced in the stored maize grain after three years, traces
of them remained after up to seven and eight years ofstorage. Filutowicz &
Bejnar (1954) observed no relation between the age of sugar beet seed and
the degree of infection by Cercospora beticola.

CI-IE.~C.'\.L EFFECfS

The importance of micro-organisms in the safe storage of seeds has

been demonstrated. Prevention of the destruction of the seeds by either
fungi or insects can be controlled by chemic:l1 treatment of the seeds. Many
new materials for such uses have become available within the last twemy.five years. For the most part, however, seeds have been treated immediately
before planting. For long-term storage o[ chemically treated seeds there
remains the question of the effect of the chemical on longevity. A considerable amount of ,york has already been done along this line.
The use of recommended methods of treating seeds ,vith organic
mercury compounds is gready restricted when the seeds are to be kept
for some years. Clayton (1931) found that treatment of vegetable seeds
with mercuric chloride, liquid organic mercurials, or -.hot- water, greatly
shortened the life of vegetable seeds--even when it caused no immediatc,
apparent injury. Dust treatments were safer. From a case of injury to
wheat germination c..msed by treatment with Cercsan, it was conduded
that under improper storage conditions treated seeds nu)' be permanently

46

OTHER FACWRS AFFECTIKG LONGEVIn'

damaged (Crosier, 1930--3). Initial seed injury caused by careless handling

is also a factor in damage by chemicals. Some fungicidal dusts, especially
those contlining ethyl mercury phosphate, have been found to depress
germination of wheat grain after storage at 20C. or higher (Wark, 1942;
Koehler & Bever, 1956). An org::mo-mercury dressing of wheat seeds
resulted in an increased percentage of abnormal seedlings when the seeds
dusted had been superficially dry, the percentage of abnormality rising to
as much as 15-3 when the seed had been superficially moist at the time of
treatment (Brett & Weston, 1941)- Srorage of treated wheat seeds in closed
containers caused rapid loss of viability, while untreated sceds under the
same storage conditions retlined a high germination capacity for morc than
a year. Over-treatment of cereal grains with the organic mercury compound, Panagen, did not appreciably impair the germination after eight
months of storage; but longer storage periods might cause the development of injury from large dosages (Andren, 1945). Fumigation of wheat
grain with various chemicals is apt to result in reduction or loss ofgermination capacity, but does not necessarily impair baking quality (Corron el 01.,

1946).
The effect of seven different mercurials on maize seeds in storage was
studied by Koehler (1938). Some injury resulted, but when the storage
was in a warm building or in a closed, unheated warehouse, the yields of
plants grown from stored treated seed were frequently as good as or even
better than those from freshly harvested seed. Treated maize seed was
definitely injured by storage under the roof of an open shed.
Kreidow & Garber (1946) stored seeds of alfalfa, red clover, Ladino
clover, and Sudan grass, after treatment with New Improved Cercsan,
Semesan. Arasan, Spergon, and Yellow Cuprocide. Open and sealed
containers were used at 10 and 2jC. Germination tests made periodically
in Petri dishes and in soil showed no injury to alfalfa, red clover, and
Ladino dover, from any of the fungicides. Sudan grass seeds were injured
by treatment with New Improved Ceresan, as shown by reduced germination within a month from treatment and by their complete loss of viability
within six months. None of the other chemicals used reduced Sudan grass
germmatlon.
Ceresan treatment, wet or dry, had no effect on the germinability of
flax-seed stored at moderate temperatures and low humidity for three-anda-half years in New ZeaJand (Black, r946).
Proper drying for at least four days, followed by treatment with
organa-mercury compounds, has been recommended as a treatment for
jute seeds (Ghosh el aI., 1951).
Polyploidy was induced in stored beet seeds by treatment with
Improved Ceresan (Lynes, 1945). After one year of storage the induced
polyploidy ranged from 0 to 17.30 per cent, and seeds stored for nm years
showed from 1'13 to 6g02 per cent induction. The occurrence of polyplaids in the field would be of little importance, for few of them would

47

I
SF.r.n PRFBERVATJON AJ\'D LONGEVITY

survive. However, a large percentage of polyploids could mean

reduced stand.

:J.

much

Conon-seed may be treated with organic mercury dusts at any time

after harvest and thereafter stored for periods of up to seventeen mon,ths

without injurious effects resulting from the treatment, and without

decreasing the beneficial effects of the treatment in increasing seedling
emergence and yield (Miles, 1939. 194I).
Sulpha drugs have been 11sed to inhibit the mould respiration in damp
grain (.Milner, 1946). Finely gTound sulphonamides, in concentrations of
from 1 :2,000 to I :250 (ratio of weight of sulphanilamide to weight of
damp seeds of wheat), wcrc dusted onto the seeds. The strongestconcenl.fation of the chemical, I :250, prevented increased respiration and resulted
in 84 per cent germination as compared v.ith 10 per cent in the controls.
Five commercial brands of fire extinguishers applied at a dosage rate of.
0'5 oz. (If gm.) per lb. on six seed stocks (LaCIU((/ s/lliva, Lolium sp.)
Triticum aestr"um, Hordeum vulgaTe, Afedimgo saliva, and Bela 't'lflgaris)
did not reduce viability (Suskin, 1953("), This is important, for fire in a
seed warehouse, or water used to quench the fire, can cause complete los.s
of valuable seed stocks.
Studies made at the Southern Regional Research Laboratory, New
Orleans, Louisiana, oyer a period of several years, have been directed
towards a chemical spray treatment of cotton-seed which would reduce
the increased respiration in storage and hence the resulting production of
heat which causes deterioration. A number of chemicals were found to be
promising (Lambou et al., 1948) "ben used on seeds containing 12 per
cent of moisture. Among these were propylene glycol dipropionate and
4,6-bis-cWorometh}TI xylene. Combinations of these two chemicals were
also effective in maintaining viability for more than si.~ months and for
reducing the free fany acids formed in the seeds during the storage period.
Seeds were sprayed with the mh:tures as-rhey were-moved by mechanical
conve)'or to large storage bins. Condon et at. (1949) found that ethylene
chlorhydrin.:md related compounds inhibited heating in moist flax- and
cotton-seeds and dCt.'Teased fatty acid formation.
In addition to mose reponed by Lambou etal. (1948) and Condon et
af. (1949), some 300 other chemicals have been tested ar the Southern
Regional Research Laboratory, and about fony-fivc-ofthese, when used in
low concentrations, have been found to reduce the amount of heating
caused by decay in conon-seed (Altschul, 1949). Contrary to other seed
treatments which arc designed to control organisms associated "ith thesecd and its deterioration, these chemicals bave tbeir effect on the seed
itself, through its metabolism. This method
been patented (Altschul
et al., 1951).
Control of infestation by" insects during storage, especially of grain
seeds, is of gr<'''at economic importance. Besides the-ntlmerous insecticide...
now on the market, D3ehthalene has becn used to control inS0ct5 on

nas

48

,
:

Phata. by eallrfeJY Department of Planl Pathalogy and Bawny, Uniursity af Minl/rsola, 51 Palll,
Minncsola, U.s.A.
PL,m ,

Sound and germ-damaged wheat seed. Doth were sIDred at 75 per eent relari,c humidity (moisture
COntent ].. '710 I49 per cent) and at 25~C. for seven months. Left, non-inoculated remained
sound. Right, inoculated "ith Aspa-gil!us raln'ctu$ became germ-d=gcd. (From Papavizas &
Christensen, 1957.)

OTIIER FAcrORS AFFEcrING LONGEVITY

vegetable seeds (S. L. Singh & Singh, 1935). No effort will be made here
to re,,;ew the literature on insect control. Recent articles by Conan et al.
(1953), and S. W. Bailer (1955, 1956) deal with this subjcct. The
physical conditions of seed storage, such as below-freezing temper::llures
and desiccation, mar prcycnt insect infestation or d~troy the insects
already present. The rice weevil, for e:xample, cannot breed in a seed with
a moisture content of 8 per cent or less, and will soon die if restricted to
such seeds for food (Conon & Frankenfeld, 1945). Other insects, such as
the flour beetles, on the contrary. are capable of breeding in very dry
seeds, but can be controlled br Io\\-lcmperature storage of the seeds. Two
interesting methods for the detection of imic:;ible insects in seeds are (i)
carbon dioxide output of the grain samples (Ho.... e & Oxlcy, 1944), and
(ii) radiographs of the seeds which disclose the insects inside (Milner et

01, 1952).

49


VI
LONGEVITY OF VEGETABLE AI\'!) FLOWER SEEDS
THE next three chapters 'will deal with the viability of seeds of specific
types under >arious conditions.
VEGETABLE SEEDS

The storage of vegerable seeds in such a way that seedling vigour as

well as viability may be retained over :l period of years, is a vital problem
to both seedsmcn and growers. As we have seen, successful storage conditions must include major consideration of temperature, moisture con-

tent, and oxygen supply. The effectiveness of the method depends on

the interrelation of these factors rather than on a single factor.
One of the earliest rcports on the subject was lint of Kunze (1881:),
who listed types of herb and vegerable seeds divided into life-spans of
from one year to eight to ten years. Those with the shortest life-span were
Coriandrum salr,;um, Sola1lum me/ongello, and Satureja !Jor/ens;s; those
which lh'ed longest were Cucumis sa/rollS, CuclIl11is 1I1elo. and Cucurbita
pepo. Carruthers (1911) stored forty-three kinds of farm seeds in paper
bags in cabinet drawers. His tests showed that the seeds varied in the
rapidity of the decline in gennination. Carrot seeds lost their vitality at a
uniform rate and gave no germination in the tenth year.
The storage of vegetable seeds under controlled conditions is compar:uivelr recent. Joseph (1921)0). working \\;th parsnip seeds, cited the
beneficial effect of low temperamre and desiccation on keeping quality,
and Beattie & Taunan (1950) kept parsnip seeds at low temperatures for
six years. Parsnip seeds lose \;ability T3pidly wben subjected to changes of
temperature and humidity (Myers. J935). Rose (1915), in his report of a
study of delayed germination in economically important seeds, remarked
that two varieties of lettuce seed improved in viability as they grew older,
at lease up to the end of the fourth year. This improvement, he said, was
due to the increased permeability of the inner sced-eoat. In his progress
report on various kinds of seeds harvcsted in 1924 and stored through
1932, Pritchard (1933) listed those of carrot, cauliflower, lerruce, and
onion as seeds which lost their genninative ability to a. great extent, while
seeds of tOrDato were a.mong those retaining their viability ycry well.
A preliminary report on the effects of different storagc conditions on
the retention of viability of some vegeroble seeds was published by Barton

;0

LO~GEVITY

OF VEGETABLE A,\''"O FLOWER SEEDS

(1935). Seeds of carrot (DIlIiCIiS carolil. L. var. saliva De.), cauliflower

(Brassica oleracea L. var. botrytis L.), eggplam'aubergine' (Soltllllllllllle!ongma L.), lettuce (Lactuca sativa L.), onion (AlliulIl cepa L.), pepper
(CapsiculJl ftutescem L. [C. all/lUum L.l), and tomato (LycopersiclIlIl esculeJltll111 Mill.) were stored, in 1932, at room temperature and at approximately -4C. Samples of each kind of seed, with the original moisture
content, were stored both open and scaled. In other samples part of
the moisture was removed, either by drying in a desiccator over calcium
oxide until about one-third of the moisture was removed, or by mixing
direetly with the amount of calcium oxide required to remove one-third
or one-half of the moisture present in the seeds. All of the dried lots
were sealed and stored at the two temperatures mentioned above. Sealed
storage was in tin cans with tight-fitting lids, sealed for the first few years
with Dekhotinsl..;r cement and later with scaling wax. Care 'was taken not
to heat the seeds in sealing.
Tests for viability were conducted both in electrically-controlled ovens
and in the greenhouse. The oven tests were made on moist filter-paper
in Petri dishes, while the greenhouse tests were made in soil composed of
peat, sand, and sod soil, mixed in equal parts. In the Petri dishes the
appearance of the hypocotyl was taken as the criterion of germination,
while, in the greenhouse, actual seedling production, or the emergence of
the shoot above ground, was recorded. This distinction is especially
necessary with seeds that have been stored for some time, as old secds
often possess enough energy to send out a short root but cannot produce
seedlings when plamed in soil. Lots of 100 seeds each ,vere used for the
germination tests. Preliminary trials at different temperatures indicated
the following to be best for oven tests of the various seeds: carrot, 15 to
30e. daily alternation; cauliflower, 25C. constant; eggplant 'aubergine',
20 to 30e. daily alternation; lettuce, 20e. constant or 15 to 30C.
daily alternation; onion, 25C. constant; pepper, 25C. constant; tomato,
2Se. constant.
Germination tests have been made on the surviving seeds each year
since their storage. Selected data obtained after up to hvcmy years of
storage are shown in Table VIII. Other records are to be found in the
original publications (Barton, 1935, 1939, 1953a). Periods of up to six
years demonstrated tomato and eggplant seeds as the most resistant to
laboratory temperature-regime storage. Lettuee and onion seeds were the
least resistant, and carrot and pepper seeds were intermediate, in their
response to this temperature regime. Further tests revealed that carrot,
eggplant 'aubergine', and tomato seeds, were still capable of about 50 per
cent germina.tion after open storage for eight years in the laboratory
(Table VIII). The life-span was extended by drying the seeds before
sealing---either by placing the seed samples over calcium oxide in a
desiccator or by mi.ung the seeds directly with weighed amounts of
calcium oxide, aftcr which the lots were placed in sealed containers.

5'

SEW PRfSER\'ATIO:\,

1\.,'0

I.Q'GEYITI'

TAJlLE VIII
GR.\IIK-\TIO"': OF nGETAUu.!:Il::'.EDS 0:-. MOISf FILITR-PAPI:.K AT CO:'\IROun>

TE-\IPERATlJRf.S AYI1:R srORAGE U:-..mR ".!RIOQi COl'\nrnoZ"S

Smngl:

e..-

un",.

Op<n

Moisture

Eggpl~nt

86"

Air-dr)'
la-I

Air-dry
10"4

,
0

,
0"...
Lettuce

5(":1lcd

Op<n
Onion

,s'

""',..
13'

Air-dry
8'2

,,

,s'

""'''''
Op<n

""'''d

content

,..=<t
0

Open
Sealed

.m-dry
12'5

,
Air-dry

6, 6,
59 6,
6, jZ

"

Sc::lled

Per cent gen:n.iJUtion :afler

Storage:l.l -4"C: for
)"C:lfS indir:::zted

"

6,

Arr-d<y

8g
9' 9'
95

...

0
0

9' 9J 6,

"

'
"
,

52

6,
8,

",

8g ,6 S,

"

.,

z:

"9'

"

"

" " "" """ "

"8,8, "8, " 8, "
"'" " " ,6 S,
80 '9

79 86

86

86
79 8, 80 S, 8,
8, 8, 8, 8, 8, 88 86

..".
.."
... ". . .
..

9'

9' 8,

95 95
9J gO

8, gO

75

0
0

,6

"

9J

95 88

gO 8g 88
95 8; 9J

"

iZ

6,

"" s, "
jZ

73 13 66
13 6, 6,

57

'9

13

6,

"'0

'0 6,

J.I

6, 6,

9' 90 ,8 86 88
9J
9' 9'
9' 8, oo 9'
90 90 '9 9' 9' oo
90 9' 8.1 90 90 9' 88

"

9' 9' 90 7J 45

9' 8g 90 86 79

"

'0 8,

.' ,

gO

So 86 66

J9 39

8g is

6, 66 6, S, 6,
68 74 6,
68 6,
6, 6, 6,
6, 73 6,
6,
66 66

gO 8, 8,

,6

9' 95 ,8 95 88 8, 8,

0
0

gO 8, 6,
gO 9J 6, JJ
gO 95 9' ,6

oo

,6

gO 8, 8, 86
95 gO
75 93 So 77 So
9'
9J ,8 9J 8, 8, ;6

9J ;> 66

, "

68

.,

,
,

., ""

""
" '" " " " ""

8, ,6 0
8, 8, 8, S,

10'0

, ,

90 13 59
86
68 J7 "
8,
'0 7J 8,
8, 66
86
S, ii
9J 88 68

"
,, " " ." ,
""

'0,

,6 '0

55 6, ,6
33 57 ;, JJ

0",.

9".,

, "

66 6, ;,
60 60 0

,6

TomalO

afrtt

in bboc:uory fOl'
)UI"!i indicated

stonge

.,. ""'' ' ,

,

c.rro.

Per cent germination

'3

.. Ptt cent gcrmin:J.tion :l.t beginning of stonge.

t At the beginning of the srorage pcriocJ.
II Dried m'er CaD to n:rno"e about onc-{hird of indie:unl moisture; b:- mi'l:etl with sufficient
CaD to rcrno\"C :l.bour onr-rhinl of lhc indicm:d moistun:; r = mi:lnl '\rim sul1kicm Cao 10 remo'"C
llboln OfIO'ohalf of me indic:lted moisrurc.
- Indicate<: tlt3r no tcst w.lS m:ldc.

5'

LONGEVITY OF VEGETABLE AI\,'D FLOWER Sf.EDS

Although some vegetable seeds remained viable for several years at

laboratory temperatures, -4C. proved far superior for keeping the seeds
(Table VIII). Even onion seeds, the germination capacity of which is
seriously impaired after only a single year in open storage in the laboratory, gave 93 per cent germination twenty years after they had been mixcd .
with calcium oxide, placed in sealed containers, and kept at -4C.
continuously_
Some vegetable seeds were stored by Pedro (1936) with and without
calcium chloride at temperatures ranging from 0 to 28C. The beneficial
effect of drying was shown significantly by all seeds after 312 days in
storage-wilh the exception of tomato seeds which 'kept' wcll in all
conditions. From his results the author concluded that dry, warm storage
was better than moist, cool storage. lIenee he considered moisture to be
more detrimental than temperature. Vegetable seeds have been kept in
Japan for ten years in air-tight sealed containers with calcium chloride
(Kondo & Kasahara, 1941).
Because onion seeds lose their viability rapidly, they have been the
subject of special investigations (Beattie & Boswell, 1939; Brison, 1942;
F. R.13rown, 1942; Myers, 1942; Nutile, 1957). All of these papers report
the advantage of drying, sealing, and low temperature, for maintaining
the viability of these seeds, and Brison (1942) pointed out l.h:J.t in room
temperature storage for forty-four months, onion seeds sealed undcr a
vacuum proved slightly better than those sealed in air, though this was not
apparent in cold storage.
The effects of moisture and temperature on the keeping-quality of
onion seeds has been studied by Beattie & Boswell (1938), who reported
that at both 4ooP. and room temperature-regime storage, 10 per cent of
moisture resulted in greater loss of germination than 6 or 8 per cent, but
was without effect at 20F. (-7C.). Seeds of high moisture contents
kept at 40F. or room temperatures showed better germination after open
than after sealed storage.
Many investigators have stored vegetable seeds under a variety of
conditions and have reported the germination behaviour of these seeds
after storage. Among such authors arc Pritchard (1933), Stone (1938),
Boswell el al. (1940), Rodrigo & Tecson (I940), E. H. Toole (1942),
E. H. Toole et a/. (1948), Harrison (1956), and Madsen (1957).
In addition to the literature dealing with vegetable seeds in general,
there are numerous reports on the keeping quality of specific types.
A comprehensive physiological and microchemical study of lettuce
seed viability has been made by Grifliths (19.J2). After two years ofstorage,
deterioration of tomato seeds was evident from 'narural' storage temperatures, but no deterioration was evident after storage at approximately
.J0op. (Myers, 1946). Be:l.ns of various kinds were the subject of investigations by Garner & Sanders (1935), Rodrigo (1939), E. H. Toole & Toole
(1953), and Dextcr et al. (1955). Data on radish (RaplwllIH sat/VIII L.) seeds
j3

SEffi PRF.sERVATIQN ANn LONGEVITY

also point to the temperature-moisture relationship in seed viability

studies (\Vaggoner. 1917 j Harrons & l'v1cLean1 1945)'
Although we do not usually think of true secus of pomco in connection
with the propagation of this vegetable, it is important to be able to store
such seeds to maintain a certain stock, or to protect hybrid seeds. Cold
storage of potato seeds at 3zoF. (0C.) was decidedly superior to storage
at room temperature, and sealing was beneficial (c. F. Oark, 1940). At
low-temperature storage, the germination of potato seeds remained
unimpaired for thirteen years and they still gave 17 per cent germination
after t\vcnty)'ears (Wollenweber, 1942). High relative humidity or sudden
changes in atmospheric conditions are thought to be detrimental TO the
viability of potato seeds-hence the value of sealing (Stevenson &
Edmundson, 1950). Similar results from storage have been obtained by
I-Iaigh (1952) and C. H. Coleman (1953), both ofwhorn report that some
potato seeds show a dormancy when freshly harvested and may not reach
full germination capacity until 4 to 8 weeks later. Okra seeds with 8 per
cent moisture content can be stored successfully for at least seven years
at 35 to 400P. (J. A. Martin & Crawford, 1954).
Deleterious effects of high moisture and temperature have been
demonstrated for carrot seeds stored for as little as ten days at 30 to
40C. and with T] per cent of moisture (-lUi & Wal:aru1be, 1955).
Thus we have seen that low moisture content and low temperatures
--especially temperatures below frcezing-are best for e..xtending the
life-span of vegetable seeds. The harmful effect of below-freezing temperatures for some seeds, however, has been pointed out by Weibull
(1955), and was discussed in Chapter IV.
FLOWER SEEDS

The rapid deterioration of many flower seeds ullder ordinary methods

of storage has led a number of seedsmen and florists to publish notes at
various times on their O'\'\'T1 personal experiences. These notes have not
usually included experimental data, but, for the most part, have simply
emphasized the need for well-planned experiments from which reliable
storage methods could be determined. llodger Seeds, Ltd., El Monte,
California, U.S.A., issued a bulletin in 1935 entitled Valuable biformation
for SeedsmCll (Anon., 1935). The bulletin lists many flower seeds in
tabular form with information on longevity and germination. No special
storage conditions are described, however. Some tests conducted by the
California Federal State Seed Laboratory, in which sixty-four samples of
flower seeds involving thirty genera were stored for Jen years, represent
extensive work to obtain necessary daL1 on the longevity of thL'SC forms.
The results obtained from this experiment have been reported by Goss
(1937) Although this report contains much information of vital importance, it does not shed any light on the possible extension of life under

54

LONGEVITY OF VEGET.WLE

.\t,u

FLOWER 5J::J::D5

special storage conditions, as all of the seeds were merely placed in paper
eme1opes and stored in a tin box in the seed laboratory at SaCTamento.

OPEN IU

SEALEO R.l:

OPEN+

SEALEO +~c

MONTHS OF' STORACE

FIG. S.-Graphs indicating \~abilit) of annual delphinium seeds of the 192';, 1925, and
1926 crops, as shown by gcnnination tests on moist fil!e:r-paper at 15C. after storage in open
or sealed containers at room tcmpct:lture (R.T.) or +8C. for different lengths of time after
Deeember 1926.

Delphinium seeds, especi:llly those of the perenni:ll forms, lose their

viability rapidly under ordinary conditions of storage. Increased interest
:in delphinium culrure has sened to emphasize this problem. Annual and
perennial delphinium seeds of the 1924-, 1925, and 1926 crops, were placed
under special storage conditions:in December 1926 (Barton, 1932, 1933,

55

SF..ED PRESERVATION AN"U LONGEVITY

193Qb, 1953a). This means that two-year-old, one-year-old, and fresh

seeds were stored. The storage temper:nures used were room temperature,
8"c., and -15C. up to 1933, after whieh the g0c. storage lots were
placed at Soc. and the below-freezing storage was at -5C. Scaling of
these seeds was effected in small glass vials with cork StOppers covered
with paraffin. The sealed vials were then placed in large bottles fitted w-ith
ground-glass stOppers.
Fig_ g shows the effect of open and sealed storage at room temperature
and g0c. on the keeping quality of annual delphinium seeds stOred for
o to 1..j.3 months. In their responses the three seed-crops showed similar
trends throughout, despite the fact that the 1924 crop seeds gave
an initial germination of only 27 per cent as compared ,nth 57 and 72
per cent for the 1925 and 1926 crops, respectively. Open storage at
room temperature proved most deleterious, causing rapid falling off in
viability until all the seeds were worthless after thirty-lITe months of
storage. When scaled containers were used, however, the decline in the
power to genmll:l.te was less rapid. Germination after sixty-two months
in scaled containers at room temperature compared f,l'murably with that
obtained after twenty-six months in open containers. Seeds in open
storage at g0c. fared slightly better than those in open storage at
room temperature but did not 'keep' as well as those scaled at room
temperature--especWlr after eleven monlhs of storage. Scaled storage
at g0c., however, served to keep these seeds viable for r43 months.
In Fig. 9 is depicted the behaviour of perennial seeds under similar
conditions. Here germination of seeds from -ISO'c. srorage has been
shov.n rather than of those from g0c., as the superiority of the lower
temperature was demonstrated early in the storage period. Also, only tbe
1925 and 1926 crops are pictured as the 192+ crop was worthless when
rccei\-ed in December 1926. Again, the slmila.r trend in beha\;our in spite
of initial germination differences is to be noted. Open storage at - TSoc.
was far superior to open or scaled storage at room temperature for preserving the germination capacity of these seeds. Sealed storage at - r SoC.
proved best of all. Here seeds which were relati\'el~T fresh when stOred in
December 1926, kept their viability unimpaired for 1+3 months.
Tests made after different periods of storage ha\'e shown that these
old delphinium seeds were not only capable of germirutting on filter paper
but also produced seedlings in soil (Barron, 1939b). Some of the seedlings
thus produced from seeds uken from the best conditions after 12-3 months
of storage were grown to maturity. The appearances of typical annual
and perennial plants are shown in Plate 4(0). These seedlings compared
favourably with those obtained from fresh seed planted at the same rime.
Seeds produced on the plants from old seeds were viable and when planted
produced norm:li seedlings. The delphinium seeds, both ::mnual and
perennial, which were stored in 1926, survived for nineteen years when
kept at temperatures below freezing (Barton, 1953Q ).

S6

LON"Gf.YITY 01' VEGETABLE Ai'\l1) FLOWER SEEDS

Deeter (1938) obtained 90 per cent germination from delphinium

seeds stored in an envelope at room temperature at Danville, Pennsylvania, U.S.A., for fivc-and-a-half years. The report of Goss (1937) included
both perennial and annual delphiniums. He found that the former
deteriorated more mpidly than the latter, though there was some difference in the individual crops. There is no doubt that the life-span of these
seeds can be increased by storage under f.wourable conditions.

Of"fN R:t

40

Z
0,"

">z

~o

>z

\
OPEN -l~C

ffi..

oO~~:::~'~O?'~~---'.'m:'o~:::~o~---~",:'---~~~~=~:::~,,,,
MONTHS OF STORAGE

FIG. 9.-Grdph~ indk:ning vilbiliry of PeTellIlEll delphinium seeds of the 1925 and 1926
crops lIS shown b~' germinatiOn 1e5{S on moi~t filter-paper at 15~C. after storage in open or
sc:llcd containers at room tcmpcraz:ure (RT.) or - '5"C. for diITl'tenl lengths of time afler
December 19:26.

Seeds of an aster (CallisfepJlus chim:nsis Nees) were stored, using three

temperatures (room, Soc., and approximately -fOC), three moisture
contents (from about 4 to 8 per cent), and open and scaled storage
(Barton, 1939b). Two types of stornge containers were employed: hermetically sealed glass tubes which were not opened from the beginning of
the experiment until the viabilit}, tests were made, and tin caos sealed
with scaling wax which were opened at the timc of each tcst and re-sealed.
To maintain viability of these seeds in open containers for periods of
longer than one-and-a-half years, it W:lS necessary, under the general
conditions of tills experiment, to use :l temperature below freezing-point.
There was some indication that deteriomtion was more rapid in open
containers in a Soc. room with a saturated atmosphere than at ordill.1ry

;7

SEED PRfSERVATiON ..\ND LONGEVITY

room temperature when the humidity was not so high. However, the rate
of decline in germination power in the two latter storage conditions was
similar, the seeds being worthless in both instances after storage periods
of longer than one-and-a-half years. When dry seeds were ..kept in sealed
containers, the length of life at both room temperature am! 5C. could be
prolonged-at the latter temperature for as long as nine years. A temperature of -4C. permitted retention of viability for at k-ast sixteen
years in open or sealed storage (Barton, 19S3(l). Verbena seeds have a
shorter life-span than aster under the above conditions.
Winter-flowering sweet pea (La/lryrUJ), pansy, Giant Golden ~een
(Viola), and VellidiUIIl seeds, were stored both air-dr)' and with reduced
moisture contents at room temperature, 5C., and approximately -4C.
(Barton, 1939b). The limited supply prevented extensive tests being made
with seeds of the sweet pca, but they remained viable under all conditions
for thirteen months, and in scaled containers for thirty-three months.
Limited tests with pansy seeds indicated that open storage in a saturated
atmosph.cre at Soc. was harmful, the seeds being much reduced in
viability after nineteen months of storage. Open storJ.ge at room temperature permitted retention of viability, though this was somewhat reduced,
up to thirteen months, when the seed supply was exhausted. Sealed.
storage at room temperature kept the seeds viable for nineteen months
but proved deleterious when the period was extended to thirty-three
months. Under the conditions of Barton's experiment, pansy seeds kept
best at 5C. in sealed containers. Ball (1935, p. 49), writing of pansy seed,
stated that 'under favorable conditions its viability will extend over two
years, but no responsible seedsman depends on this'. He further remarked
that storage conditions must be favourable in order to keep the seeds in
good shape until the August after harvesting, but he did not describe
these favourable conditions. Harrold (1935) believed that some of the
failures attributed to poor pansy seeds were really caused by high soil
temperatures or a hot dry atmosphere, pointing again to the importance
of adequate testing procedures. Goss (1937), reporting on the results of
storing pansy seeds in paper envelopes, presented data which showed
45 per cent germination after five years of storage and 2S per cent after
nine years. The results of other tests (Barton, 1939b) also indicated that
pansy seeds may be kept viable for periods of longer than two years, and
probably their life-span may be increased several-fold by sealing seeds
with approximately S per cent moisture for storage at a low temperature.
Venidiu//l seeds of poor quality maintained their vitality for thirtyseven months under a variety of storage conditions (Barton, 1939b).
Over a ten-year period, Heit (rgS7) tested the germination responses
of seeds of forty-two types of flowers in the laboratory, greenhouse, or
field, and concluded that germination failures arc not always caused by
weak or old seeds, as adverse weather and soil conditions, poor cultural
practices, insects, and diseases, are all important factors in survh-al.
58

LONGEVITY OF VEGETA1II, Ai\.'D FLOWER SEWS

It h:J.s been generally recommended th:J.t lily seeds be planted while

they are fresh. Craig (1931), however, stated that Lilium auratltm will
start better from year-old seeds than from fresh seeds. He also found that
Lilium regale seeds wruch had bcen stored for five years at 'Veymouth,
Massachusetts, germinated nearly 100 per ccnt. Oement (1938) reported
morc rapid loss of viability of L. regale seeds. He obtained about 80 per
cent germination from 1936 and 1937 crops planted on 12 April 1938,
but he secured only 8 per cent germination from the 1935 crop treated
in the same way. Thus he found that these seeds kept for n"O years bur
were practically worthless after three years. Neither Clement nor Craig
uescribeu the storage conditions employed. It is presumed, however, that
open storage under ordinary room conditions of temperature and humidity
was used.
Seeds of the regal lily (Lilium regale Wilson) have been kept under
various storage conditions at the Boyce Thompson Institute for fifteen
years, germination tests being made at least once a year (Barron, 1939c,
1948), Mter completion of preliminary germination tests, moisture
determinations were made prcparatoJ}' to storage. Air-dry' seeds were
found to contain 99 per cent moisture, calculated on the basis of the
dry-weight of the seeds. One lot of seeds was placed in a desiccator over
calcium oxide for two days, after wruch they were stored. Determinations
made on this second lot showed 4'5 per cent moisture. Seeds with this
reduced moisture content as well as air-dry seeds were stored in scaled
containers in the laboratory, at 5C., and at approximately - S"c. Air-dry
seeds were also stored in open containers. Sealing was effected by using
tin cans with tight-fitting lids scaled with sealing W:1.."(. Tests for determining viability were made in moist granulated peat-moss at a daily
alternating temperature of IS" to 30"e., and in soil in the greenhouse.
In the case of the alternating temperature, the cultures were kept at J5 "e.
for sixteen hours and at 30"C. for eight hours each day. Germination at
the controlled alternating temperature was practically complete after two
weeks, while a slightly longer period-up to twenty days-was required
for the completion of seedling production in the greenhouse.
As there was very good agreement between germination at controlled
temperature and seedling production in the greenhouse, dara for the
latter only are presented. Also, results of viability tests after J, 2, 30.5, 7,
I J, 13, and IS years were selected as representative of deterioration under
the various storage conditions. These data are shown in Table IX.
It will be noted that air-dry seeds remained fully viable in the laboratOJ}' for one year but were reduced in germination capacity after nyo years,
none bcing viable after three years. Sealing had no appreciable effect on
the seeds under these conditions. However, when part of the moisture
was removed anu the seeds were placed in sC'J.led containers, good
germination could still be obtained after five years of storage in the
laboratory.

59

<

SEED PRESERVATION Al"\rn LONGEVIn'

At Soc., open storage proved unsatisfactory for keeping seeds ,iablc.

No tl."Sts were made up to the three-year period but at this time the
gcnnination capacity was so reduced that only 16 per cent produced
seedlings. Sealing was effective in keeping seeds with 9-9 per ccnt moisrure
viable at Soc. for eight years, but thereafter there was a gradual decline
in germination ability to 8 per cent at the cnd of fifteen years. With reduced moisture content the longevity at Soc. was C'.\tcnded.

TABLE IX
REG..iU. LILY. SEIDLI:>:G PRODUCTIOS IN SoIL [N TIlE GREE1\'HOUSE ..u'TER
STOR...GE. FRESH SEWS GAVE 86 PER CE\i SEEDLL'i"G PRODl'crIO~

Stonge condirions
Moislure

Tempc:r.l_ content Op<nM

rurn
percentubor:ltory

5"C.

9.

-S"C.

4'5

".

.".

4"5

Approx.

""'"
0

4"S

S
S
S
S

Per cent seedling production iller stor.agc

No. of rears indicated

,
88

'7

.'

.p
35

9'

..

s.- '7
,

s.-
oS

"
'7

.3 s.-

" "
0

'7

'7

", '3

"

'7

ob

86

,.

31
80

88

'3

"

'7

.' . ..' .'.'

'5

rOT

"" .,,.
"
'5
s.- .3

C..aIcub!ed on ~ b2si5 of the dry-weight of the seeds.

-

lndiatcs th2t no test was m.o.de.

The effectiveness of Soc. as a storage temperature is thus demonstrated. The cause of the rapid loss of viability of seeds stored open 3.t this
temperature is to be found in the high humidity prevailing in this room.
The combination of this lower temperature and high atmospheric
humidity proved only slightly superior to the higher temperature but
lower humidity of the laboratory for keeping regal lily seeds. "then rhe
absorption of excess moisture was prevented by the use of air-right
containers, seeds remained viable for at least eight years at Soc. When
this lower temperature was llsed for storage, scaled air-dry seeds ("ith
9'9 pcr cent moisture) kept well but, at the higher temperature of thc
laboratory, it was necessary to reduce the moisture content in order to
permit retention of vitality for longer than two years (Table D...').
At -Soc. both open and sealed storage kept the seeds viable for
fifteen years. There may have been an indication of reduced germinationcapacity of the seeds from open storage at the fifteen-rear tcst (74 pcr cent
seedling production), but this remains for future tests to confirm.
60

LONGEVITY OF VEGETABLE Al\'D FLOWER SEIDS

"-ithin the limits of tills experiment, the moisture content of the sceds
stored at a temperature below freezing-point had little or no effect on
their keeping quality. As storage periods lengthen, the superiority of
below-frcezing temperatures over Soc. for maintaining the viability of
many k--inrls of seeds is being demonsl:r.lted. It should be poimed Ollt,
however, rhat approximately ~ Soc. is the lowest temperature used in
these tcsts. Other experiments will be necessary to determine the effeets
of still lower temperatures.
Plate 4(b)A shows the storage temperature effects as related to open
and scaled containers, while Plate 4(b)ll shows the effect of reduction of
moisture content on sealed storage in the laboratory, at Soc. and -Soc.
In sealed storage at ~ Soc., these seeds of regal lily remained fully viable
for fifteen years (Barton, 19S3a).
The results reported here are in agreement with those obtained by
Clement (1938), who probably had storage conditions in Ontario similar
to the open laboratory at Yonkers. A drier atmosphere could have
accounted for the extension of the life of rcgallily seeds to five years as
reported by Craig (1931).
In conclusion it may be said that if a dry cold-storage room is available,
seeds of the regal lily can be kept fully viable for at least ten years. If the
atmospheric humidity of the room is high, it is necessary to seal the seeds
(which should be thorougWy air-dry, i.e. contain less than IO per cent of
moisture) unless a temperature below free:t.ing-point can be used. If no
cold-storage chamber is available, the seeds may be kept at room temperature for five ycars by reducing their moisture content to approximately
.5 per cent and placing them in sealed conrainers.
Germination tests of air-dry Gladiolus seeds stored under controlled
conditions have shown that they can be stored successfully for twelve
years at ~4 c. l or at Soc. if the container is not opened in the interim
(Barton, 195312-)
The viability of Gmtian12- crillita Froel. seeds was determined after
storage under various conditions (Gicrsbach, 1937). Secds stored in open
containers at room temperarure and Soc. showed a complete loss of
viability after one year. Seeds sealed in air at room temperature had also
lost their germination capacity by this time, whereas those sealed in a
partial vacuum at room temperarure and those sealed in air or a partial
vacuum at Soc. still retained their original germination power.
Knudson (I9oP) was able to store seeds ofCattleya for fourtecn years
and still obtain germination, but seeds of other orchids could not be kept
so long. The same general conclusions concerning orchid seeds were
reached by Svihla & Osterman (I9H), but they found further that these
minute seeds could be frozen and dehydrated and still remain vi.able.
Seeds of the tuberous begonia and snapdragon, which are two to ten times
the size of orchid seeds, were similarly treated but failed to sur\':ive.
In our enthusiasm for storage at low temperatures we should not lose

6.

SF.ED PRESf.RVATION .l,ND LONGEVITY

sight of the possibility of damage to some seeds. Weibull (1955) conducted

experiments on the storage of flower seeds from 1952 to 1955 and found
that they showed very definite reactions to storage at -20C. He classified
the seeds studied into four groups, as follows:
was favourable for storage of Kochia scoparia, Myoso/is
sylvatica, and Callistephus chinmsll.
2. -20C. seemed favourable, but the results were less clear, for Dalllia
variobilis, Gypsophila pouiculata, Primula malacoidcs, and Zinnia
elega1ts.
3 No difference between _woe. and 'ordinary storage' was found for
Chrysanthemum spectabile, De/phinium IlybridulIJ, D. ajacis. Verhmo
veJlosa, and Salvia sp/mdms.
4- -20C. seemed to be disastrous for AlltirrlJiJ1111ll mojus, Asparagus
pIUI1l0S11S, Petunia IrybTida, and Viola tricolor vaT. ma.riml~.
T. -20C.

It is possible that this e.xperiment was not continued long enough for the
superiority of the below-free'Ling temperature to become evident in
e:ttegorics 2 and 3. as it often is not manifested until the seeds have been
in storage for five years or more (Barton, 1953 a).

62

VII

LONGEVITY OF SEEDS OF FIELD CROPS

THE keeping quality of .wheat grain is of great interest, not only because
good seeds are vital to good crops, but also because of the food value of
the grain. Conditions which keep the seed good for planting purposes also
maintain good food value and taste.
Carruthers (19II) \"as onc of the first to record the germination of
wheat seeds of different ages. He tested so-called white and red wheat
seeds of the 1896 crop each year for nine years. Fresh seeds gave 100 and
99 per cent germination, respectively. After five years of storage these
percentages decreased to 88 and 80. Extension of the storage period to
eight years caused a further reduction in viability, so that only 29 and 51
pcr cent of white and red wheat seeds, respectively, wete still alive at the
end of this time. J. White (1909) recorded 2 per cent germination of a
South Australian whcat seed-lot which was sixtccn-and-a-half years old.
Although these early records indicate that wheat has a short-lived seed,
later controlled e."'periments have sho\\11 that the life-span can be extended
materially by storage under controlled conditions. One of the first of these
to be instigated was that ofWo)"lI1per & Bradley (1934), who stored fresh
seeds of Rivet wheat at room temperature in sealed tubes under a variety
of conditions. No germination tests were made on these seeds until 1927,
fifteen-and-a-half years after the commencement of the experiment, when
only one lot was found to be viable. This lot was of seeds placed in tubes
with dry calcium chloride.
Further storage was made of seeds of two wheat varieties after drying
in a desiccator over calcium chloride (\Vhymper & Bradley, J934).
Desiccation of the variety 'Square Heads Master' to 4.10 per cent moisture,
followed by scaling and storage in the laboratory in the dark, preserved
viability for up to about nineteen years. Highly desiccated Rivet wheat seed
(moisture content of 066 per cent) still showed germination of 82 per
cent at the end of fifteen years and nine months of storage. In some of the
experiments air 'was excluded by covering the seeds with wax, immersion
in oil, or sealing in an atmosphere free from oxygen. The authors state
that it appears from the results that these methods caused cessation of
respiratory exchange, resulting in death of the seeds, whereas restriction
of respiration by desiccation extended viability by the retardation of
63 .

SU.J) PRFSERVATION AJ'\,'D LO:\'GEVITY

intracellular reactions. Toxic substances may have accumulated in the

seeds when gaseous exchange was restricted. The aurhors predicted that
the life of wheat seeds may, under the ba.1: conditions they used) be
approximately fifty years. A later n:lcase on the performance of these
same seeds after twenty to thirty-two years of storage showed a decline
in germination from 82 per cent after twenty years to 69 per cent after
thirty-two yeaTS (\Vhymper & Bradley, T947). Under the experimental
conditions used, wheat seed with a moisture content of 43 to 4.8 pcr cent
dropped in germinative power 17 per cent in nineteen years and 31 per
cent in thirty-onc years, or about I per cent per year. At this rate, the
authors point out, their estimate of fifty years for the life-span of the seeds
is likely to he too low. Unfortunately, the seed supply has been exha.usted
and so no more tests can be made.
D. W. RoberSOn and co-workers in a series of publications (D. W.
Robertson & Lute, 1933, 1937; D. W. Robertson et at., 1939, 1943)
emphasized the importance of humidity on wheat seed longevity. They
found that length of life increased as the humidity W3..<; decreased. In an
unheated room in Colorado, germination decreased 7 per cent in the first
ten years of storage, and by the end of twenty-one Tears only 128 per
cent germination was obtained. These seeds contained 95 to 114 per
cem of moisture as compared v,.;th only 43 to 4.8 pcr cent in the seeds
stored by '\/hymper & Bradley. The highcr the moisture content, the
shorter was the life-span and the greater the injury from exposure to high
temperatures (Whymper & Bradley, 193-, 1934b).
Swanson (1941) has also demonstrated the relation bern-eell temperature and moisture content, and recommends that high-moisture wheat
grain be stored at a low temperature. ''''heat seeds were placed in 4-oz.,
stoppered hottles (I) in the laboratory and (2) at 41 OF. Germination after
laboratory storage of seeds containing 10 per cent of moisture was 92 per
cent after eighteen weeks of storage. Increasing the moisture content to
14 per cent decreased the germination to 51 per cent after eighteen weeks
of storage, and a further increase to 16 pcr cent of moisture resulted in
the death of all the seeds after only nine weeks. On the other hand, the
germination capacity remained unaltered upon storage for up to eighteen
weeks at +loF.-regardlcss of the moisture content, which was frOllLlo to
20 per cent.
In 1938 wheat grain was stored in Japan at ~25 to -3S c C., for one
year only (Tejima & Misonoo, 1938). Viability was good at the cnd of that
time but no further report is available. In the light of other favourable
responses to below-freezing temperatures, these wheat seeds should have
a long life-span.
In addition to loss of viabilit)" high moisture content of wheat seed
causes heating when the grain is stored in large lots; it also inereases mould
growth and brings about deterioration of baking qualities of the flour
(C. H. Bailey, 1917, 19I7a; IHcCalIa ~t at., 1939; Lafferty, 19.P; Anon.,
64

(01) Delphinium plants produced from seeds stored for 123 momhs before planting.
plants; 1'1, perennial plants.

A,

Annual

')

PI.<\TE 4(b) DeYelopment from regal lily seeds sown in flats after four and a half years of storage.
.1, Flats sown with air--dry 5ei:ds stored in the hooratory (/4i), at 2joC. (Ulltrr), and at -j0c.

(right): (I) in open containers, and (2) in sealed containern. II, flats sown with seeds dried OVer
CJ.lciurn oxide before storage at \llriOU5 tempcrarurcs: (I) in the laboratory, (2) at Soc., and
.
(3) at - Soc.

PI..An: 5

Pi"ux tada: sccdlin~ production in rhe greenhOllse aft~r storage fur fi,"c years (air-dry seeds).
~,Planted in flat difL"t:tlr from stora"" tlasks: (1) open at mom temperature in dark, (2) ~ealed at
room temper-nure ill <.lark, (3) in ,"aCl1um al room temperature in dark, (4) in ncuum at room
tcmpttal>.= in light, (~) Optll at .5~c., (6) srnl~d at S'c., (7) in '"<leu.urn a~ 5 c., (S) ':ren in
refnger:lllon room (- 5 LO -15 C.). B, Planted In fbt after One momh III mum granulate peatmoss at 5 C. Numbcrs as in A.

:
:

f.ONGF.t'ITY OF SF.F.DS OF FlaD CROPS

I :5; E. P. Carter & Young, 1945; Milner ~t al., 1947, 1947a; Harris &
\hlsrcr. 1953; Oxley & I1yde, 1955). Storage conditions maintaining
fu.I1 \-;abilitr also retain desirable food characteristics.

lJe\-elopments in the growing, harvesting, and marketing of malze

there known as 'corn') in the United States necessitates a car'ry-o\er of
~ surpluses of gr<lin from one year to the next as a means of price and
supply stabilization. Also, the special techniques, time, and labour
an lIved in the production of the homozygous inbred lincs necessary for
subsequent production of superior hybrid maize, make it imperative that
;ood storage conditions for maintaining viability of breeding-seed stocks
%S well as planting material be known. Most of the litcrature on maize is
f American origin, because America. is the greatest 'corn' producer.
Argentina is also important in world production, but the maize grown there
is of the 'flint' type and, because of the dry autumn climate, dries down to
moisture limits that arc safe for storage and shipment. Tn other parts of
the world where maize seed is grown, it is either dried carefully by the
indi\;dual grower who has a small supply, or it is consumed ,,;thout the
:1tteSSity of storing.
In a book by Huelsen (1954), a general account of the taxonomy,
morphology, and physiology of sweet corn will be found. He devotcs a
dupter to the factors, including StOragc, affecting germination and
'=l'Owth of the seed.
- Detailed experiments with ~arrow Grain Evcrgreen sweet com stored
under different conditions of temperature and humidity have been
ttpOrted by Boswell el al. (1940)..Moisture and germination testS were
lD3de at intervals of ten days for up to 251 days of storage. The original
;ermination capacity of the seeds was 82 per cent. Estimated germination
\'"31ues are given, based on the trend of the curve for successive tcsts.
torage at 80F. in humidities of 78,66, and 44 per cent resulted in IS,
and 70 per cent germination, respectively, after storage for 110 da:rs.
Seeds stored for a similar period at 50F. and in 81, 66, and SI per cent
humidity germinated 6S, 70, and 72 per cent, respectively. No germination
""as obtained after storage for the fu11251 days at the high humidity and
temperature. At low humidity and temperature significant deterioration
lnS evident, but the seeds still gave 72 per cent germination after 251
d3.ys. The short total time of storage prevented the appearance ofdeterioration under all storage conditions, bur favourable moisture and temperature
safe storage for short periods are indicated.
Yellow Dent maize seeds of the 1921, 1922, 1923, and t924 crops were
s:mred in an unheated room in Colorado (D. W. RobertsOn et al., 1943).
Samples were taken for germination in February of each succeeding
E

65

SEED PRESERVATION AND LONGEVITY

)'tar. Viability gradually declined [rom the (irst to the tWl'nty-.first year,
the seed.s germinating J2 per cent at the end of I..he paiuu.
A morc detailcd test was started by Sayre (1947) in 1933. St:ed lll:tize
with the moisture content adjusted to 5, 75, II, 14.6, or 182 per cent
was placed in large glass culture tubes. The tubes were then stored at
86, 72, 37-39, 16"_28, or oF. In 1938, another series was starteu,
using tin cans for storage. Germination tests were lll:lde each )'e:lf for
each series. All of the seeds with the highl,ost moiSture content of 18'2 per
cent which were stored at room temperaLure lost the power to germinate
before the end ofone year. Seeds were more tolerant to the higher moisture
contents as the storage temperature was lo\\ered. Thc SWlC author .had
shown previously (Sayre, 1940) that low temperatures (3 and -25C.)
and absence of a:..'Ygen from the storage container extended the life of
maize seeds.
Confirmation of the roles of humidity and temperature in determining
the keeping quality of maize seed is available in the works ofDuvcl (1909),
Brunson (1946), S,",-aine (1957), and many others. Drying seeu maize willI
electricity (Duffec, 1937) or with different drying agents (Larter, 19+7)
before storage has been found effective.
The yield of high quality open-pollinated maize declined gradually
after the second year of storage, but still retained satisfactor}' viability
and vigour after five years (Dungan, 1940). Limited tests made by Dungan
indicated that some hybrid sced maize produces a better yield than others
after onc ycar of storage.
Haber (1950) also demonstrated that thcrc is a genetic basis for longevity of seeds of sweet corn, while confirming that humidity and temper;lturc are the major environmental factors responsible for maintenance of
germination capacity. His n-perimental results showed that, in general,
the longevity of hybrid seeds was better than that of the two inbreds used
to produce the single-cross hybrid. When two inbreds with good longevily
of secds were crossed, the hybrids had the same character. This was also
true of inbreds 'with short life-spans. Long-lived inbredxshort~lived
inbred hybrid seeds rerained viability much longer than the short-lived
parent. It would be necessary to store inbreds with a short-lived seed
character under filVourable, colltfollcd storage conditions in order to
maintain such stock.
Enough 'work has been done on seeds of maize and of other plants to
prove the value of below-freezing storage for long life (see Chapter 1V).
Advantage is being taken of this knowlcdge to prepare cold-storage rooms
at about oF. for the maintenance of valuable maize seed stocks (Anon.,
1949, .1950). l",lcRostie (1939) made a study of the thermal death-point
of maize from low temperatures (down to -10F.). Seeds with moisture
contelHs above IS per cent suffered severe damage which was greater
under fluctuating than under constant temperatures. Safc storage (5f maize
grain requires a moisture content well below TS per cent.

66

J,

LONGEVITY OF SEEDS OF FIElD CROPS

While the food value of maize grain may be directly rdatcu to its
viability, the popping quality of popcorn i1ppears to he entirely indepcnJcnt
of germination capacity (Stewart, 1936). Ability to pop is retaincJ at least
fourteen years--si.'\': years beyond the ability of the particular lot of popcorn to germinate.
OrnER CEREALS

The life-span of cereal grains has been reported by various authors

a... follows: wheat, 6 to 32 years; oats, 6 to 29 YCilrs; barley, 2 to 10 years;
rye, 9 to 10 years (Burgerstein, 1895; Todaro, 1898; Carruthers, 1911;
Sifton, 1920; Percival, 1921; Welton, 1921; D. W. Robertson & Lute,
f933j Walster, 1943; Haferkamp, 1949). An atmosphere devoid of
oxygen, or, alternatively, drying and sealing, extends the life ofcereal grains
(Bussard, 1935). Drying is imperative if losses from fungal invasion are to
be avoided in combine-harvested barley grain (MacLeod, 1952). The
malting industry has to sell, for feeding purposes, many hundreds of
bushels of barley grain annually because of the loss in storage of gerruinabiliry. The greatest damage to malting barley grain occurs during the
early storage period before the grain dries out. Storage conditions in the
State ofWashingron are exceptionally favourable for retention of viability
by cereal grains (Haferkamp et at., 1953). Moisture content and respiration arc important in the heating and deterioration of oat seed (Bakke &
Noecker, 1933).
General discussions of the physiology and biochemistry of grain in
storage are given by D. W. Robertson et af. (1939), Slusansehi (1939),
Kretovich (1945), and Blum (1954). Causes of deterioration in, and
practices for storage of, cereal grain are discussed by Oxley (1950a, 1955).
Anderson & Alcock (1954), in the detailed book they edited on the storage
of cereal grains and their products, state in the preface that millions of
tons of cereals are wasted every year through spoilage of various sorts in
spite of recent advances in technology.

GRASSF.S
Che,",ing's fescue (Pes/ilea rubra L. var. COlm/llllala Gaud.) seeds have
claimed the attention of several workers bcr:ause of their economic
importance and their short life-span under certain storage conditions.
Foy (1934) stated that deterioration of these seeds in transit was due to
excessive heat and humidity in the hold of the ship. Viability could be
maintained by lowering the moisture to about 5 per cent, or by shipping in
cool storage. Seeds stored over calcium chloride in the laboratory for seven
years still gave over 90 per cent germination, while undried seeds (about
13 per cent moisture) were dead in two-and-a-half years. Commercial
drying could hcst be done by heat. Twenty-one samples of Chewing's

67

SEED PRESERVATION Al\'D LONGEVITY

fescue seed stored in the laboratory for from twenty-one to thirty-four

months lost 6 to 69 per cent of their original viability (Patrick, 193 6).
Kearns & Toole (t939(1) found that the advantages of shipping these
seeds under cool conditions from New Zealand were lost within a few
months after arrival in the United States, unless cool storage was continued
or the seeds were dried out. Seeds dried before shipment and kept dry
after arrival in the United States gave germination percentages after
eighteen to twenty-six months' storage at room temperature which were
equal to those of non-dried seed on arrival. Tests were made using domestic
seed and storing it in sealed containers at _10", 2",10",20, and 30"e,'
after adjustment of the moisture content to 8 to 14 per cent. Here again
the interrelationship of temperature and moisture was shown. The
higher the temperature of storage, the lower was the moisture content
required for maintaining viability. When the seeds are stored at 30ce,
the moisture content should not exceed 8 per cent; at 20"e it should
not exceed 10 per cent; at lOoe it should not exceed 12 per cent.
FJuctuation in moisture content is injurious to Chewing's fescue secds
in storage (LaPine & Milberg, 1948). Gane (I9481i) found that the usc of
artificial atmospheres of nitrogen or nitrogen+ I per cent oxygen had no
noticeable efrect on the rate of deterioration, while lowering the storage
temperature and reducing the water content of the seeds increased their
retention of viability during a four-year storage period. Kearns & Toole
(T939) emphasized the need for low temperature in the germination testing
of Chewing's fescue seed. especially when the seed is fresh.
As early as 1894, Samek published a table showing the results of
germination tests of twenty different forage plants for a period of cleven
years. The founeen species of grasses, etc., included in the study showed
decreased germination after five years, the actual amount varyi.ng with
the species, and all of them produced only occasional seedlings after ten
years of storage. The grasses, etc., reported on were: French oat b'T:lSS
(Avella etatior), Italian ryegrass (Lolium italieum), English ryegrass (Lolium
perel11le), tall fescue (Fes/llca pmtemfs), sweet vernal-grass (Antlzoxanthum
adoratum), meadow foxtail (A/apecufIls pra/rnsis), timothy (Phleum
pr(lfeme), orchard grass (Darty/is g/omerata), blue--grass (Foa pratetzsis),
crested dogstail (Cynosurus crisllitus), fiorin (Agrostis stolonifera), sheep's
fescue (Fest{(ca ovilla), hair-grass Desdzampsia (Arra) ftexuosa, and spurry

(Spagula. arvf1l5is).
Using hulled and unhulled seeds of timothy (Phlcum prate1lse) stored
in six different warchouses in various districts of Sweden, Esbo (195+)
found that unhulled seeds maintained good germination capacity until at
least the beginning of the second summer. Hulled seeds declined more
rapidly, especially as a result of the tempemture fluctuations of the first
summer.
Excess moisture should be removed from grass seeds as soon as
possible aftcr harvcsting, according to Williams (1938).

68

LONGEVITY OF StlJ)S Of F1ELJ) CROPS

It has been observed that certain native grasses in Austmlia can

disappear for a while owing w drought or animal injury or grazing, and
then reappear (Myers, 1940). Seeds of some of these grasses were kept in
packets and stored in the laboratory at Sydney under ordinary armospheric
cunditions for twelve years. Panicum antidotale gave 3 per cent germimtion after such storage for twelve years, but 88 per cent after seven ye-drs;
P. de:compositum gave 3 per cent gennination after nine years, but 63 per
cent after four years; P. pro/uium gave 2 per cent germination aftcr eight
years, but 65 per cent after four years; PaspaIidium jubifIorum gave only I
pcr cent germination after ten years, but 72 per cent after five years; Dalttbonium semiallnularis gave 5 per cent germination after four years, but
100 per cent after one year. The grcatest gcrminatiun in every case was
from seeds one or more years old, thus indicating a dormancy in freshly
harvested seeds.
Seeds of Andropog01l !urcaftls, A. scopan'us, Sorghastrum nu/ans, and
Boute/oua curtipendu/a, are also dormant when fresh and do not begin
normal germination until from fOllrteen to eighteen months after harvest
(Coukos, 1944). After dormancy has been broken by storage under dry
conditions, germination remains at a high level for several months. Cold
storage in scaled jars extends the life-span to at least thirty-eight monthsexcept in the case of Sorghastrum nutans seeds, which remained viable
for only twenty-four months.
Italian r:{egrass seed could be stored sllccessfully for two years if the
seeds were dried to 13 per cent of moisture (Gorman & Greenwood, 1951).
Longer storage pcriods required further reduction in moisture content.
One crop of carpet-graK'i (Aronopus affinis) seed remained fully viable
for three years of storage while the germination of another crop fell to 5
per cent in three years. The difference may have been due to trearment of
the seeds before they were received for special storage (E. H. Toole &
Toole, 1939).
For general information on the different phases of grassland cultivation, the reader is referred to a recent book entitled Grassland Seeds
(Wheeler & Hill, 1957).
LEGUMFS

Because of their importance as forage crops, alfalfa or lucerne (Mcdicago satrva L.), various clovers (TrifO/ium spp.), and sweet-clovers
(Meli/otus spp.), have been thesubjects ofstorage invcstigations. Terasvuori
(I930) obtained some germination of seeds of different species of TrifOlium
,vhich had been stored for fifteen to thirty-four )7curs. Stevens (1935)
found that the viability of seeds of alfalfa and sweet-clover stored in the
laboratory decreased from about 95 per cem to 60 per cem, and of red
clover to Ie pcr cent, in twenty years. !vlost of the impermeable seeds of
alfalfa became permeable during the first year, but a few remained

69

SEED PRESERVATION At'iD LONGEVITY

impermeable after twenty years. Clover seeds became permeable more

slowly, and 'hard' seeds persisted for the longest time in sweet-dover.
Later reports by Stevens (1943, 1954) included germination data from
six samples of alfalfa and two samples each of sweet-dovcr and red dover
seeds stared in ordinary seed envelopes in ventilated metal boxes in the
laboratory. Alfalfa sced germinated 37 to 70 per cent after thirty years and
showcd another 10 pcr cent decrease at a total age of fOIty years. All
samples of clover seed deteriorated more rapidly, giving only 1 to 13 per
cent germination after thirty years. Myers (I952) stated that if lucerne
seeds arc to be kept in a dry climate, they can be in either open or sealed
containers, but to remain viable in a moist climate they must be sealed.
The persistence of viability of seeds of alsike clover after fifteen years
of storage was attributed to the 'hard' seeds in the sample (Lafferty, 193 I).
We have already seen (Chapters I and IV) that the presence of an
impermeable COat extends the life-span of many seeds. Similarly, the loss
of viability in storage may be caused by a softening of the coats. There are
often changes in coat permeability and hence germination capacity
between harvesr and the time for sowing the following year, as has been
found for red clover seeds U. N. Martin, I945).
Some leguminous seeds have been claimed to show signs of rencwed
vigour after having beh'1lll to deteriorate (Rodrigo, I935). This may be
related to failure to treat the seed-eoats for germination or to seed-coat
permeability changes during storage. The date of harvesting determines
the moisture content and hence the suitability of seeds of velvet-beans
(Stizolobiufll spp.) for storage Oustice & Whitehead, I942).
Seeds of blue lupine (LI/pinlls fillgustijoJillS L.) containing 20 per cent
of moisture can be stored without damage for long periods only at relative
humidities of less than 70 per cent, and probably only at ones of less than
65 per cent (Brewer & Butt, 1950). High quality, naturally cured seeds of
less than IO per cent moisture content can, however, be stored without
great damage for eight weeks at a relative humidity as high as 85 per cent.
These seeds were injured by drying artificially to 3.6 per cent moisture, so
that their germination was reduced from 85 to 38 per cent, but tillS low
viability was maintained for eight weeks even at 100 per cent relative
humidity.
Seventeen different crop seeds were tested by Sonavne (1934) over a
period of twelve years. In general, leguminous seeds retained viability
longest. The storage offarm seeds by Vibar & Rodrigo (1929) demonstrated
the advantage of sealed containers in maintaining viability.
Storage of the important k'gume, soya bcan, will be discussed in the
next chapter.

:
:

VIII
LONGEVITY OF OTHER ECONOMICALLY IMPORTANT
SEEDS
OILY SEWS

(Glycine l1lu,lJ\'lerr.)
In spite of the importance of soya bean production and the relatively
short life-span of the seeds under ordinary conditions, detailed work on
rhe requirements for successful storage has been done only recently.
Oathout (1928) was one of the first to report that high temperature and
lack of ventilation caused rapid loss of viability in soya bean seeds with
moisture contents abo\'e 14 per cent. J\'Icasurcment of the damage to
soya bean seeds under the ordinary conclitions of temperature and
humidity obtaining in Trinidad (monthly maximum from 83 to 87F.,
monthly minimum from 65 to 72F.; humidity at the saturation point
almost every- night) was made by G. E. L. Spencer (1931). Germination
tests of samples stored at 55 to 60F. were also made. After cool storage
for ten months, roo per cent germination was obtained, while the percentage was reduced to II after storage for the same period under ordinary
conditions.
Soya bean seeds of eight different varieties, which had been treated
,,~th an insecticide, were stored in North Carolina for periods of up to
four years and five months (Burgess, 1938). At tlle end of the storage
period, germination varied from 7-1- per cent for the Biloxi variety to 0 ptr
cent for the Haverlandt (or Herman). Seeds of all varieties germinated
[rom 95 to 100 per cent at the beginning of the test, so varietal characteristics are evidently among the factors determining life-span.
Increasing the moisture content from 9'4 to 19' I per cent caused rapid
Jas...;: ofl'iability ofsoya bean seeds stored at room temperature in Minnesota
bmstad & Geddes, 19-1-2). Seeds kept in storage for two years or longer
in lllinois gave poor fidd-Sl::l.nds (Burlison el al., 1940). Ottawa, Ontario,
Canada., apparentl)' affords a better climate for soya bean seed storage;
seeds not more than three years old germinated satisfactorily, though not
quite as well as fresh seeds (LaugbJand & Laughland, 1939). Four-year-otd
Sttds showcd COlL"iderable wcakness and anything older was unfit for

SOYA BEAN

pbnting.
A morc comprehensive experiment was starrcd in 1934 to determine

7'

5F..F.D PRESERVATION

.~',i1)

LONGHTfY

the effects of temperanlre of storage and moisture content of soya bL':ln

seeds on their life-span (E.l-L Toole & Toole, 1946). Seeds of the soya bL':ln
varieties Otootan and Mammoth Ycllow were adjusted to moisture comenL';
of approximately 18, 13, 9, and .5 per cent, and phccd in sealed gbss pint
fruit-jars at controlled temperatures of _10, 2. 10, 20", and 30"C.
Germination at the time of storage was 93 to 97 per cent. TesLS, m:ide :it
intervals after storage, indicated a similar response of the two varieties.
The seeds \\;th the highcst moisture content lost viability rapidl~' at the
higher temperatures bur maintained good germination for 6\'e to SL\:
}""t.':lrs at -10C. Air-d.ry seeds (approximately 13 per cent moisture) fell
rapidly in germination capacity at 20" and 30"C.; there was, howevet. no
loss until after three years at IOC., and the original viability was retained
to the end ofthe experimcnt at _TOC. Seeds ",ith approximately 9 per
cent of moisture showcd slight deterioration after a year at 30"C.. but
r:l.pidJy lost germination power after that time. Lo\\'ering the temperature
to 10", 2, or - IOe., maintained full viability for ten years of seeds with
9 per cent of moisture. DT)ing the seeds to., per cent moisture content
caused immediate injury a.c; e\;denccd by cracks across the cot)'ledons,
sC'o'crance of \"llSCldar connections of some cotyledons, and severance of
the plumule.
Other evidences of the deleterious effect of excess moisturc on SO\':l
bean seed viability, together with additional data on longevity, haye ~
presented by Akamine & Rippenon (1938), Ramstad (1942). Germ (19.n)J
Tcn'ct (1913), Robbins & Poner (1946), Milner & Thompson (1954), and
a series of articles by Milner & Geddes (1945. 1945", 19-1-6, 1 946a). Some
of these authors also deal ,,;th respiration, heating, and moulds induced
by excess moisture.
COliON

(Gossypium spp.)

Regions where cotton is grown arc usually humid wilh high tempcra-

tures-eondirions kno'i\" to be detrimental to most seeds. Cottunseeds

are no exception. For example, in Charleston, South Carolina, t"Ottonseeds held in bab"S deteriorated rapidly aftcr two years; but those in which
the moisture content had been reduced below 8 per cent, and which had
been stored in tin containers, showed only a slight decrease in viability
after seven years, and some of these seeds were still gcrminable after ten

years (Simpson, 1942). In a series of publications, Simpson has given dau

on the conon-seed storage problem, induding the results of a controlled
storage experiment begun in 1937 (Simpson & Stone, 1935; Simpson,
1935, 1940, 1942, 1946,1953, J953a).
In the controlled experiment, corron-seeds of two upland varieties
were adjusted 10 levels of moisture from 7 to J 4 pcr cent and storctl in
sealed containers at constant temperatures of <)00, 70, and 33F. Corresponding controls were subjected to normal air temperatures at Knoxville,
TeIlllessce. At 90F., seeds ";th 14 per cent moisture were all dead in

72

::

::I

~_.

;;;

LONGEVITY OF OTHF.R ECO!'OMIC.....LLY IMPORTANt SEEDS

fuur months, and those with reduced moisture were either dead or nearly
so within three Ye'Jrs. All seeds stored at 33F. for three years 'kept'
without appreciable deterioration, but those with 14 per cem moisture
began to deteriorate after five years at this low temperature. At lower
moisture contents, seeus at 33F. maintained viability after seven years.
Seeds stored at 70F. were intermediate in response between those at 90
and those at 33F. Seeds at 33 and 70F. were continued in storage for
up to fifteen years, when the driest lot (7 per cent moisture) still gave 73
per cent germination (after being at 70F.) and 91 per cent germination
(after being at 33F). Seeds with J3 per cent moisture which had been
stored at 33F. were also still viable after fifteen years (72 per cent), but
those with 14 per cent moishue content were all dead by then. Simpson
states (Simpson, 1953(/, p. 391): 'The much longer life under storage at
33F. plus the striking differences in longevity obtained for moisture
levels within and between the other storage temperatures, furnish conclusive proof of the inter-relationship of moisture and temperature in
cotton seed deterioration. The mOisture and temperature tolerances
determined in this experiment are sufficiently definite to serve as a guide
in safely storing cotton seed. If the indicated conditions of moisture and
temperature are maintained, good seed may be stored for fifteen or more
years with reasonable assurance that viability will be retained.'
In work carried out at the Hellenic Cotton Research Institute, Sindos,
Greece, Christidis (1954) used seed of the same genetic constitution
produced in successive years but all tested during the same year. Three
varieties of upland cotton served as source material. The experiment
showed that the genetic constitution remained unchanged in the individual
eascs, although there was variation in their behaviotlf, owing not only to
the age of the seed and the error of sampling but also to the conditions
under which the seeds were produced. In spite of these variations,
however, seeds of all tluee varieties germinated satisfactorily after eleven
years of storage.
Whether controlled storage conditions must bc used for cotton-seed
depends, as for many other seeds, upon the locality of storage. Thus it is
neccssary to dry the seeds and store in sealed containers at the Central
Experiment Station, Bureau of Plant Industry, in Manila, Philippines
(Flores, 1938). Sun-drying before storage is effective. A consistent
decrease in germination capacity was found in cotton-seeds stored under
warehouse conditions in California, so that the decline was significant in
tluee years (Towers & Harrison, 1949). An investigation of the causes of
deterioration of conon-seed in the equatorial province of the Sudan
(Anthony & TaIT, 1952) showed that loss of germination capacity was
closely related to the relative humidity. The optimum humidity for storage
was found to be about 33 per cent. This was much better than higher
humidities and somewhat better than a dry atmosphere, which indicates
that excess drying is injurious. Phillis & Mason (1945) also found that

73

SF.f.D PRESF:RV.o\1l0N .""'ill LONtJr:VITY

therc was a surprising loss in viability of cotton-seeds stored under

conditions of vcry high aridity. Ho\\cvcr, if seeds werc transferrcd from
o to 50 per cent relative humidity for si.-'<. months before they were germinated, their performance was improved over that produced by planting
the seeds directly from the low humidity storage_ The failure to germinatc
after extrcme desiccation is apparently not due to the development of
impermeable seeds, as such seeds swelled readily \yhen moiStencd and
some of thcm produced short radicles. Howcver, these seeds died \\;thollt
furthcr dcvelopmcnt. Artificial drying of cotton-seed at the gins cncndcd
the life-span and prc\-entcd the dcvelopment of ~ fatty acids (Rusca &

Gordes, 19f2).
Formation of frce fatty acids in stored seeds, and the usc of chemicals
for desiccation and exrens:ion of life of couon-seed, arc discussed in
Chapter V. Infecrion by bacteria or fungi \\as not associated "ith loss of
liability in tests conducted by Arndt (1l).l6).
FL.o\."\

(Linum spp.)

Moisture and temperature ha\-e again been the factors most studied
in connection with this and other oily seeds. In North DakQ[2, fla..,--st:eds
from seven to fourteen years old gavc satisfactory germination but
samples from fifteen to eighteen years old showed loss of viability
germinating from 50 to 89 per cent (Dillman & Toole. J937). Germination
of fresh seeds depended upon the particular harrest. A wet Mrvest and
threshing season caused darmge to the seeds. Also, seeds of high moisture
contcnt (from 10 to 18 per cem) wcre injured. Flax--set"d of good quality,
stored under favourable conditions, could be used for seeding for a period
of six or eight years.
Decker & Reitz (19+8) stored Linom flax-seed of high germinating
power at 40F., about 7ZoF., and 86'T'., after adjusting the moisture
content to three different levels (9, I I, and 13 per cent). Storage was for a
period of 940 days, with periodic sampling for laboratory and field
planLing:s. Conclusions from the da.ra obtained were: that flax-seed cannot
be stored safely at B6F. at moisture levels of 9 per cent or above; that
moisture contents above II per cent caused rapid deterioration after
ninety days in storage at 72F. j and that a tcmperature of 40F. 'was best,
regardless of the moisture content.
High moisture COntent and injured secd-eoats influence the oil
(]uality of flax-seeds (Painter & Nesbitt, 1943; KopcIkovskii e/ fll., 19;(J).
P F~o\NUTS

(Arachis Jrypogaea L.)

Well-eured peanut seeds held in the shell under room tempera lure
condiLions kept well for at least five years and the quality '\25 not improved
by storage at 32 or 40F. (Beattie, 1931). Shelling the seed before storagc
caused slightly greater decline in germination capacity. Similar results

74

LONGEVIlY OF OTl-IF.R ECONO,\UCALLY 11I.lPORTA."'IT SEEDS

were reported later by Beattie el al. (1932). Excess drying before shelling
increases kernel breakage (Beattie & Kushmao. 1947). Mathur et ai. (1956)
recommend tbat shelled peanut seeds should be stored at 3Zo to 35~"
and unshelled ones at room tcmpcramre. G. E. L. Spencer (1931)
reported a small but significant difference, in f<lYOur of the cool storage,
in the germination of seeds held in cool and ordinary storage.
C-.'I.."TOR OiL

(Ricinlls sp.)

The quality of seeds of Ricinus declines rather rapidly during the first
few months after harvest and then more slowly. so that 75 per cent of their
original vitality is still present after tcn years (Guillcmet, 1931).
OTHER SEEDS

Whether maintenance of the viability of tobacco (Nic()tirll/{~ sp.) seeds

is:l problem also depends upon the place of storage. It was reported carly
in this century (Shamel & Cobey, 1907) that thoroughly dried tobacco
seeds could be kept with perfect safety for ten years or even longcr.
Similar results were obtained in Wisconsin U. Johnson et al., 1930) and
in Connecticut (Anon., 1936). In places like Florida and Puerto Rico,
however, seeds more than two years old arc apt to produce inferior stands.
Poptzoff (1933) recommended 55 to 60 per cent relative humidity and
a temperarure not greater than 15C. for the storage of tobacco seed.
According to Poptzoff, excess drying does not injure the seeds, but rather
prolongs their life-span. Immature seeds do not keep as well as ripe ones,
which should be stored at low relative humidity and a low temperature
above freezing-point (Chirkovskii, 1938).
Kineaid (1943) controlled the moisture content of tobacco seeds by
storing them over chemicals which gave an atmosphere of about T, 40, 60,
and 100 per cent relative humidity-<lf. in some cases, seeds were sealed
in glass tubes aner their moisture content had been adjusted. Test lots
were kept in the refrigerator at about 5"C., and in a basement, a laboratory,
and an attic. After cleven years, seeds stored over calcium chloride (I per
cent relative humidity) at each location, and all of the seeds in the rcfrigerator, germinated welL Samples under normal, uncontrolled conditions
in the laboratory were nearly all dead after three years. Seeds with 5'3 per
cent or less of moisture, which had been stored in stoppered vials, had
deteriorated markedly after eight years of storage-indicating a harmful
effect of excessive drying which was not found by Poptzoff(1933)
Twenty-year-old seeds of tobacco stored dry, in the absence of air and
light, were still fully viable (Sehlocsing & Lerom, 1943)'
Sugar beet seeds are significantly reduced in germinating power after
storage for five years (E. W. Schmidt, H:I34) as shown by germination
percentages and the poor root systems of resulting seedlings, According
to Fitutowicz & Bejnar (1954) the limit of safe storage of these seeds could

7;

SEED PRJ::SERV.o\T10N .,,1\'0 LONGEVITY

be extended to seven years (for elc\'cn varieties harvested for SCHn

successive years) without decreasing their planting value.
Mesquite (ProJopis vdllti,ta WOOL) seeds may remain \;ahle for
forty-four years owing principally to impermeability of the coats (5. C.

Martin, (948).

f'

I
I

Seeds of coltsfoot (TuJsi/ago far/ora L.), which ordinarily lose their

,-iability "ithin four months at any temperature above oc., kept it for
more than two years at -15C. (Dorph-Petersen, 1928a).
Seeds of HibisClIs cannabis L lose their Ylability quickly--evcn under
the most favourable storage conditions, according to Poprso,- (1929).
Moisture content is important in the safe keeping of sorghum seed
(Shedd & Walkden, 1947). but these seeds will maintain 50 per cent of
their original viability in certain localities without special stoT:l.gc (Karpel'
& Jones, 1936).
Seeds of prickly-pears (Opun/ia spp.) germinated after forty-four
weeks, including twelve weeks of storage and thirty-rwo weeks under
conditions suitable for germination (Myers, 1939). Later tests indicated
that Opunlia seeds may remain viable for years (Myers, 1939).
Arnica mOil/alia L. seeds maintain their vlability for a year in dry
storage (Esdorn. 1940). Poor quality Arnica seeds require three weeks
for germination, while good ones germinate in two weeks.
Information is available on a few horticultural seeds not dealt ,,;th in
other pam of this book. A.mong these are seeds of Nandino domesrica,
which can be stored dry at :l low temperature for ren months "ithout
appreciable loss of viability (Afanasiev, H}4-3). Testing of these seeds is
complicated by their delayed germination which results from a rudimentary, undifferentiated embryo.
Limited tcsts with the seeds of Angelica archange/ica indicated that
scaling in an air-tight container stored at about 35cF. kept the seeds viable
for eight months, at which time the experiment was terminated (Taylor,
1948). These seeds also required a special germination temperature of
4z oP. for sixteen hours daily and of 77F. for the remaining eight hours.
Data are available for Kocltia indica seeds which indicate that they retain
their germination capacity after storage for various periods of time when
open to the air in the laboratory. Conditions tried include: open at 30"C.,
open at 38C., and scaled over calcium chloride at 30C. (El-Shishiny,
1953) No deterioration took place for more than fourteen months at 30C.
uver calcium chloride. The author states that rapid deterioration of sced
in open storage at laboratory temperatures and at 30cC. i>hows that loss
of viability is accelerated by moisture more than by tcmpcrarurc. No
doubt the life-span could have been further e.xtended by low-temperature
storage of the dry seeds.
The relaled Arabian 'Shok' (Prosopis suphamana) also has \'cry h1.rd impermeable sccd~oats. Dormancy of the seeds is readily broken by filing Ot tteatment loI-ith
concentrated sulphuric acid. :!.S was recently de.moI1s[r.ucd by Or. A. K. Khudairi
of [he Deparunenr of Doowy, Universiry of .Ba:ghrlad, Iraq.-Ed.

7tl

LONGEVITY OF OTHER ECONOMICUl.Y IMPORTANT SHillS

Blueberr)' seeds ,,'cre cleaned in :l Waring Blendar and stored dry in

vials with cork stoppers in an ordinary refrigerator at about 40"F.
(Darrow & Scott, 1954). Good g;ermination was secured from one twelverear-old lot and from two nil1c-year--Dld lots stored in this way_ No other
special storage conditions were used.
There is a widespread belief that strawberry seeds are very short-lived
hut thc~' still gave 81 pCf cent germination after two years of storage in
screw-top j:us under normal conditions for Sydney, Australia, with a
mean temperature ranging from 54F. in July to 7IogoF. in January, and
mean relative humidities of 65 to 74 per cent for the same periods
(Myers, 19S4).

77

IX

LONGEVITY OF TREE SEEDS

FOREST trees, espcci:illy conifers, arc sporadic in their seed production
(Wakeley, 1931, and others). The quantity of the harvest, and the intervals
of time between seed-bearing, both vary with the different species and
with the climatic comlicions-inc1uding the case or difficulty of pollination at the time of opening of the inflorescence. Such circumst:lllCCS make
it imperative to use stornge conditions which will ensure a constant ::."Upply
of seeds for planting.
Co~lfERS

A considerable amount of work has been done on the viability of

coniferous secd$. CieslaI' (1897) found tlmt sealed storage of Pic~a txulsa,
Pinus Qwtriaca. and Pinus syh>estrulengthcned the life of the seed. The
difference in gennination of seeds from scaled and open storage after six
years was found to be as high as 33 per cem. Haack (1909) confirmed some
of the results of CieslaI'. He found tb:1t sealed as against open storage
reduced the fall in germination power of pine seeds by 16 to 68 per cent in
different samples in three years. He also found that the seeds must be
dried thoroughly if they are to be stOf<..-d at moder:Hcly high temper:l.tures.
Drying, however] had no effect on sealed storage in a cellar or an unheated
rOOIll, but open storage was possible only in a dry room. Haack also
emphasized the necessity of a distinction bcnvecn germinating percentage
and the percentage of actual plant production. As the ability to geuninatc
on a synthetic substrate rose, the power to produce plants in soil rose even
more rapidly. This effect became more marked when the seeds were
of poor qualit)" or when seedling bed conditions were unfavourable.
Only quick-growing seeds produced plants whieh SUf'\.;wd in the seed-bed.
It was reported as early as 1880 (Sargent, 1880) that ten-year-old seeds
of PillIIS ctmtorta would germinate provided they had remained in the cone.
Coker (1909) referred to this remarkable retention of viability in still
unopened cones of Pinus attenuata and some othcr forms, and the prescm:e
. of good seeds in twenty- to thirty-ycar-old cones. He further pointed out
that the conditions existing in serolinous cones arc almost ideal for the
preservation of seed viability because of the regulation of gas exch:mge
and humidity. Additional proof of tlus was givcn by Dlumer (1910).
Tillotson (1921), working on Pillus patTin-om, P. mtmtieola, P. slroblls,

78

LOl\.GEVlTY OF TREE. SEWS

P. CollfOrro., Picco. CIIgclm01/11i, and PseudotSllga la:rifll/ill, spread out all the
c;.ecds thinly on a 1I0or and fanned them stca.dil) fur two days with an
electric [n, aftcr which he stored them untIer various conditions. Hc also
found that storage in air-tight containers wa.'i far superior to any other
form. In this condition seeds "'ere little affected by temperaturc. However,
contrary to the findings of other workers, be reported the poorest results
from low-temperature storage.
In 1928 an anonymous articlc (Anon., 1928) on the keeping-quality of
noble fir seed (Abies nobilis) reported the beneficial effect of cold storage.
These seeds were well dried before sealing in air-tight glass jars. Wakeley
(193W) stored longleaf pine seeds in tight containers at low temperatures
and found that thC)' germinated '\ ell one or two years after collection.
Seeds of 'Sequoia gigamea' were stored effectively though with decreasing
viability for eighteen years in an unsealed Mason jar at rOom temperature
(Toumey, 1930). On the other hand, the same worker reported lillie or no
germination of Ta:rodium distidwtn after dry open storage for onc winter.
Bates (H)30) stored white pine seeds for one year in containers sealed
with paraffin. He found that seeds stored at a low temperature, especially
if they had been dried previously over sulphuric acid (30 per cent
relative humidity), gave a higher germination percentage in less time from
planting than did corresponding samples of fresh seed. The low moismrc
content was equally beneficial under all temperature conditions. Seeds of
Pinus IOllgifolia whicb had been sealed or mixed with charcoal in a gunny
s:rck still gave good germination after two years (Champion, H)30).
However, the best rcsulLS were obt:a.ined when the seeds were well dried
and sealed before the damp season. High moisture content and variable
temperatures were found by Coile (1934) to cause a loss of vitality in
slash pine (Pinus caribaea Morelet) seeds.
Isaac (!934) demonstrated that not only did cold storage preserve
noble fir seed for five years without appreciable loss of viability, but it
actually seemed to increase its germinating power. The same author
(Isaac, 1935) showed that douglas fir seed, ifleft on the forest fioor, either
germinated or decayed within a year after it fell. Hence special storage is
necessary. Baldwin (J934) reported the early loss of viability also of red
spruce seeds when they were stored in duff, while during air-tight storage
the germination tlecreased about 10 per cent each year for the first three
years.
A det:rilcd description of the results of storing sOme conifer secds
under controlled conditions bas been given by Barton (r93511.). The
object of the experiments was to determine the effects of sealing,
temperature, vacuum, and drying, on the keeping-qualityof these seeds.
Scaling was performed in glass distillation flasks. Incase the air was to
be exhausted from the flask, the top was scaled, after which the side tube
was drawn out to a small bore and attached to an oil exhaust pump. When
a good partial vacnum had been obtained, the side tube was sealed while

79

SUD PRESERVATION

_"",,n

LONGEVIn

still all:ached to the exhaust pump. In some tests, drying: W:lS aecomplishe{i
by placing lhe seeds in desiccarors over calcium oxide for six:ty-one days
priur lu storage; in othcrs, varying amounts uf quicklime" ere-.rni.'\':ed with
the seeds before sealing. Storage was at room temperature, S"c., and in a.
refrigeration room with an average temperature of -IS<Jc. for the firsthvo or three years of the experiments, after which the seeds were trans~
ferred to another refrigeration room -with an average temperature of
- 5 to -4C. Storage at room temperature was tried in the light as well
as in the dark.
When viability tests were made, one seed sample \yas always planted
direclly.in soil in the greenhousc whilc another sample was mix.ed with
moist granulated peat-moss and placed at Soc. Samples from this lowtemperature treatment (stratification-see Glossary~ were planted in the
greenhouse after one or two, and, in a few cases, afler three, J;Ilonths. In
all cases, actual seedling production in soil was taken as :lll... index of
viability. Pretreatment at low temperatures for after-ripening was essential
for some ofche species studied. For example,.in the tests of Pinus tar:dfLL.,
if sampk-s were planted directly in the greenhouse from the storage flask
one would conclude that the viability was lost after three years of storage,
whereas with a pretreatment of one or rn'o months in moist granu1ucd
peat-moss at Soc., a high percentagc of sound scet.llings was produced.
This is illustrated in Plate 5, where.scedling production after storage for five
years is depicted. Pinus caribaea MoreIet and Pillus pOttdn-osa Douglas
showed the same marked effect of pretreatment at low temperature, while
other furills studied, though less affected, benefited from the treatment.
The seedling production data from these experiments (Barton, 19350 )
showed that sealed storage at low temperatures (SoC. or -4" to - ISC.)
was effective for the maintenance of viability. PilJus lacda seeds under
scaled, low-temperarurc storage- retained their seedling-producing
power fully for seven rears, wherea.s in open room-temperature storage
there was a decided decline in vitality after one year., and only a few
seedlings were obtained thereafter. As compared with Pinus tar:da, P.
cari/mea" P. echinata lVlili., and P. resillosa Air. all 'kept' only slighrlj'
better in open storage at room temperature and exhibited che same
beneficial effects of sealing at low temperatures. Pinus pa/us/ris .Mill. seeds
lost viability much more rapidly. Seeds from open storage :1.t room
temperature lost their germination power completely in one year. Even
under sealed, low-temperature storage there was a gradual decrease iIL
seedling production to 50 per cent of the origiml after Jive years. Pinus
ponderosa, Piaa excelsa Link., and Picea glauca (Moench) Voss (P.
calladellSis of authors), stored only in refrigeration rOOms with temperatures of -4 to -15C., all kept weIl for four to SL'\: years.
All seeds were thoroughly air-dried before the experiments wer.e
started. Artificial desiccation, whether moderate or excessive, was found
ineffective or harmful. A vacuum proved favourable to retention of

80

"
"

=
=

WNGEVITY OF TREE SEEDS

TABLE X

ru:DLlNG PRODUL,IO:\' IN TIlE ti"REE,"'lIOUSE FRO;"I PINE Sf.WS SfORED

AIR-DRY l];\,l)ER VARIOUS CO;-"'UITIONS

Stomgc
~recies" Temp.,

0c.

Per cent seedlin!,: production after storage for No. of )calS imliCllcd
Planted directly fWnl
stor:tgc

Open CO)
or sealed

(5)
8

Lob.

P.

5
uhiTlilla

-,
Lah.

" "
S\-ac. 3+
" " "
0
" '7
S-<iir " '7
3'

P.

liUJa

.." "

'0

0
0
8 ,S
'0 ;0 ,8

'5

60

S-vac. +5

,.

';

'3

"

S-Y:lC. 16

"

" "

;1 ,

$-<lir
S-'=3 1

,-,

'9
3 '7

5 '3

0
3
6 '3

"

'7 '9

60

"

35

30

I.'

"2;

,6
0

,6

6, ,6

7'

35
73

'9 33

'5

49

" "3
"

'3 ,8

'5
OS 7'

74

At the bcginninfr of the stDr:IgC experiment; P. c&hi1fatc :md

secdlin['; proUllction in the greenhouse.
- Indicatcs Ihat no test was made.

50 ,8

73

66 8g

S-'ac. 55

"3

" "
""
"
90
,8
"
78 " 60 "
;' "
"1'. tacJa " "
49

"

,
"
8, " "
" ,8 ,.8, "8," " ""
"

... "

,6 39

57

-, """0 "2; ,, "" ,, ,., ,

"
Q

'5 '0

'3

8; 77 '3 6,

'9

" ,.

55

"

"

3; ,6 39
7Z 53 67

"

,.

53

3' '3 '3 '9 ,8 ,6 37

8 '7
33 '7 " '7 "
S_\-:l.c. 52 5'
30
39 53 3'

s.-:.'r
0

,.

Pretreated at 5~C.
foc 1 month

lij

87

0
3

8g

!f.'\C up to S.j. per cent

TABLE Xl
SEEDliNG

STORED

THE

LN

PRODUCTION

GRfEl'H-IOliSE

FROM

CONIFER

SEFJ)5

LN SEALED CONT.>\J!'ir:RS AT APPROXn1.>\.ITLY --l-0c.

Per cent seedling produCtion after storage for lears

Specics

Wt. of Sft:<Is

Plamed direcrly from

I CaO. gm.

SIOr:lgtl

, ,
0 0
0 ,
0 ,

100+ 0
100+IC

P'.r.:JS taeda

'00+'7
100+:15

IOO+ 3~
100+-1-5
P!.;lJ C:::UJSIZ

Pi.~a glaull1

50+ 0
8

5+
z.;+

25+
2';+

,
0

" " "

, 0 0
0

0
0
0
0

5' 57 37
33
'3

'"

3' 5' 3'

" '50

",

,
,

0
0
0

0
0

5
0

"
3

'5
3

'5

'"" ""6, "'3

"'7,0 3', '""0
,

Pretreated at 5eC.
I month

33

,",6 "6, " " ,

"50

'0

'"

"5 ",

"

37

'3

", '"

55

3'

59

5' ,6

"

45

",

3' '0 57

3' '9 66 39 5'

6, ,6 '7 53 5' 58 34
53

'0
37

r"

56 33 5'

"
79

70 53
'9

""

,6

"

9
0

'0
,
"
, "0 0

80 8,

, , '"
63
3 6

0
"
"
.. At the beginning of the slOrage experiment Pinus lada, Pian ~.ralsa, and Pian glaucn gave
Z5+ 1 5

up '" 52, 65, and 'Ill per cent seedling production, respocth-cly.
~ Indicatcs tbat no lest
m:Jde.

8,

,
SEf.I)

PRESt:RVATION A..~ LONGEVITY

viabiliL\' when Lhc seeds were stored at room tempe1.lture, and hoth
vacuum and !ow-temperatun; storage oven;ame in part the injurious effects
of drying.
Some of these conifer seeds were tested further, after various periods
of stor.Lge, until they were no longer capable of germin:ltilln or until the
seed supply was exhausted (Barroll, 1953a). The data in Tables X andXI
will suffice to show the trend of results secured from all of these seeds. A
few effects are outstanding. Of the three temperatures tried, that of the
laboratory resulted in more rapid deterioration than 5 cc., which btter
temperature was, in turn, less favourable than ~4cC. for seeds of P.
eehitlata and P. taeda (Table X).
Seeds stored in reduced air pressure obtained by means of a vacuum
pump (v:lc.) kept better than those sealed in air or left without sealing.
This was especially to be seen in bboratory storage where some seeds in
evacuated flasks usually maintained viabiliq' for up to at least fifteen
years of storage. This is in contrast to open or air-scaled storage at this
temperature, where very few seeds survived for even two or tlu:ee years.
The data in Table X also point to the auv;mtage of pretreating rhese pine
seeds in moist granulated peat-moss for one month prior to planting in
soil in the greenhouse-an effect alreadr noted above. This is especially
marked for P. taeda seeds.
Mixing pine seeds with different amounts of calcium oxide to att:lin
different moisture contents for storage has thc advantage of being a vcry
simple and effective procedure, although care must be taken to avoid
excess drying. Seeds of P. taeda so stored for eight to fifteen years in a
room at approximately -4"C. showed the definite ill-effects of too much
dr}ing: (Table Xl). Again, the necessity for pretreatment of the seeds for
seedling production in the soil .in the greenhouse is demonstrated. Seeds
of P. paltlslris mixed with small amounts of calcium oxide retained their
germination capacity for nine years at -4"C.
Picea exec/sa and P. g/ollca can be kepr in sealed containers at approximately -4"C. for as long as fiftcen ycars pro...' dcd they arc not dried too
much (Table XI). It will be noted that the germination of neithcr of these
forms was dependent upon pretreatment in a moist medium at 5C. On the
contrary, such pretreatment tended to decrease germination in somc cases.
The efficacy of below-freezing temperatures for storage of conifer
seeds has not been generally known until recently. A statistical analysis by
Wakeley (1945) of Barton's 1935 data on pine seeds showed belowfreezing temperarures to be significantly better for storage than Scc.
Results ofsubsequent tcsts on these seeds and many others have confirmed
and extended this conclusion, as indicated elsewhere in this book (Chapter
IV). From 2 to 4"C. has heen recommended for all coniferous secd by
Heir & Eliason (1940). Noble fir seed can be held for five years at 15F.
(Isaac, 1934.), and a temperature of 32 to 39P. is recommended for white
pine secd storage (Roc, 1948).

82

LONGEVITY OF TREE SF:IDS

Many other workers have srndicd the effect.. of storage conditions and

ageing un the germination capacity of coniferous seeds, and for further

r
--=

information on particular species the reader is referred to the follo\\'ing:

Anon., 19;.1; Barner & Dalskov, 1955; Crossley, 1955; eurlis, 1955;
Goksin, 1942; Huss, 1954; Kjaer, I950; Kondo & Kasahara, 1944;
Lipkin, 1927; ,Mirav, 1946; ~'I. L. Nelson, 1940; Rchackova, H}54;
Schubert, 195h 1955; Shirasawa & Koyama, 1915; Tourney, 1921;
Vincent, 1929, 1937, I937a, 1938; \1Fuson, 1953- Additional information
nJ.:l}' be obuined from the book on forest tree seeds by Baldwin (lg.pJ
Dent (19-+7) bas written a seed manual for Indian forest plants, in which
stor.lge effec.:ts :ue included.
a\1S

Seeds of the American elm. (Ulmus amer;ama L.) belong to a group

which norm:illy liyc anI)' a short time in open-air stornge. It has been
commonly supposed that these seeds must be planted as soon as they
rmture in e:t.r1y summer. with consequent impairment of seedling stands
by the drought and heat of summer and, later, the necessity of protecting
the tender young plants with the beginning of cold weather in the fall.
C. E. Steinbauer & Steinbauer (1932) desiccated elm seeds over dif[erent concentrations of sulphuric acid for a period of two weeks, after
which they placed them in sealed containers at 0". 10", and 20"C. They
found that low temperature and low water-eontent were favourable [or
the retention of vimury. the seeds remaining 'good' for up to nine monrns
of storage but degenerating rapidly thereafter. Germination was much
reduced after eleven months, which is the approximatc time that fruits
would rove to be stored in order [Q be available for nursery planting in
many localities. The germination tests of these authors were made
between moistened blotters at 20"C. No soil tests were reported. They also
noted that elm seeds havc a resting period which is regardless of most
storage conditions but is influenced both by the moisture content of the
seeds and by stonge temperature. No actual moisture determinations were
made.
Delayed germination in American elm seeds has also been reponed
by Rudolf (I937), who obtained new seedlings in May 1937, from seeds
pbnted in June 1936. He :lttributed this del:ly to extraordinary meteorological f:Ietors which :llso made possible the retention of vitality for one
VC:lr.
. George (1937) stored American elm seeds in containers with tightlining covers placed in an attic of an unheated building. Out-door
temperatures ranged from IIS Q to -28 c F. during the period of the test.
As he obtained 79 per cent germination on moist blotters at room
temperature after ten-and-a-half months of storage, he concluded that no
special storage condition was necessary. Soil tests of these seeds yielded
33 per cent seedling production.

---~SEED PRESF.RH.TJON Al\:TJ

LO~GE\tlTY

l\'bll:l.icsi (1935) found that the germinati\'e capacity of the English

elm (Ulmus campestris L.) remained f:Lirly conSCl.nt for twO months :md
then diminished so that practically all of the seeds were dead after six
months. He also srntcd that many of the fresh seeds did not germinate
promptly but were more or less retarded up to a maximum of twenty-two

days. This recudation, he said, depended on the 'embryonic post:::

maturation'.
Some of the results obtained by these various: workers strongly
suggested that measurement of the viability of elm seeds could be
influenced markedly by the methods used for germination. This was
confirmed by Barton (1931)<'1), who tested the germination of fn.:sh seeds
of the American elm, Ulmus dm~ricana L., under a variety of conditions.
As a result the following testing procedures were used to determine the
germination capacity of American elm seeds after storage. At t3ch tesrjn~
period, fruits were mixed ",ith moist granulated peat-moss and pheed at
controlled daily alternation of temperature of 10" to 2j"C. Germination
l,;Ollnts were made when the radicle had appeared and had begun to elongate.
In addition to the oven tests, fruits were planted in soil in the greenhouse,
using a m.i.xture of equal parts of sand, sod soil, and granulated peat-moss.
Fruit samples were planted directly from the storage containers. Other
samples were placed in bottles in water and allowed to soak for twentyfour: hours exposed to light, the source of which was a sixty-watt .t\l1azeb.
lamp placed at a distance of one foot above the bottles. Mter soaking, the
water was drained off and the seeds were planted in soil. Still other
samples were mixed with moist granulated peat-moss and placed at a
eonstmt temperature of Soc. for one month prior to planting in tlle gn..' ellhouse. This method was sho\T-n to improve the seedling production
markedl)', although elm seeds are not commonly supposed to rcquirelowtemperarure pretreatment. Seedling production was recorded when the
seedlings had appeared abov~ the soil.
These methods were used to test the \;ability of American elm seeds
stored for various periods at room temperarurc, Scc., or - Soc., in open
containers or in sealed containers with air, oxygen, or a partial vacuum.
Seed moisrure contents of 7 (air-dr)"), 3, and 2 per cent were used. Results
after sixtccn months of storage are given in a paper published in 1939
(Banon, 19394 The comparative germination capacitiC'S of fruits wim
8 per cent moisture stored in open containers and in sealed rubes containing air, together with storage temperature effect,>, are shown in lTig. 1.0.
The advantage of scaling and of low temperarure is apparent. It W:1S
further shoVoIl that seeds with a reduced moisture content-were superior in
germination capacity after storage in sealed containers at room temperature for si.."{teen months. An atmosphere of ol:ygen proved espttially
deleterious when both moisrure content :md storage temperature were
relatively high. A vacuum served to prolong \;ability in cases in whieh
other storage conditions \\cre unfavourable.

84

LONGEVlTY OF TREE SEEDS

..o~--~-,---,----------,------------,

<'

-5C

,5 ,,, ,",,,
o

~
zO<>

'

',~_----1

"""

tl

--:: .----::J

"~~~--~_:'----===----- -------....---,,~

----0...

1:'1;

40

............
................

,\,

' ..........+.,.
,-

\\

01

...

~-, ... "

.... ....

~':-~-~-~-~-~-=-~-~-~-~:~ ~-:= ; ;-~';-~'~-~'~-~-; ;-J

......

~-

-!-__
.6...
4

+5C (0 - - - - - - - - -

MONTHS Of STORAGE

FIG. TO.-Graph indicating effect of stOntgc of Ameron elm seed;; under various conditions upon gl:rminarion at 100 ro 25C. d:lllr :Utern.:lrion. Dotted lines indicae open storage;
solid lines indicate !>ealed sto....ge. R T. ~ room temperature.

Up to 1939, sixteen months was the maximum life-span reportcu for

American elm seeds. L. P. V. Johnson (1946) obtained some germination
after one year of storage. Barton (19530), continuing the tests on the seeds
described above, found some of them still capablc of germination after
fifteen years of storage, as is shown in Table XlI. Again in this table is
shown the necessity for pretreatment of stored seeds to obtain a true index
of their germin!1rion capacity. A comparison of the seedling production
obtained in the greenhouse v;rithout pretreatment and after pretreatment
for one month makes this clear.
in moist granulated peat-moss at
Air-dry seeds, i.e. containing 7 per cent moisture at the time of storage,
were stored in open as well as in sealed containers in the laboratory and
at 5C., but arc nOt shown in the table because they did not survive two
fared much
years of such storage. Seeds in open containers at - 4
better, germinating up to 51 per cent after SL": years of storage. Seeds
dried to 3 or 2 per cent moisture remained viable longer than air-dry ones,
regardless of temperature. As to the effect of temperature itself, there is no
doubt as to the superiority of -4C. over laboratory or 5C. for maintaining viability. Safe storage under laboratory conditions did not exceed two

sec.

--

=E

--~

0c.

8;

I~

SEED PRESERVATION .o\..ND LONGEVITY

years. At Soc. the seeds could be stored for at least six years, and at _4C.
the life-span was extended to fifteen rears. This extension of the life of
elm seeds by controlled storage conditions is p::micuI3rly noteworthy in
view of their reponed short life-span.

--

TABLE XII
SUDLlNG PRODUCITON IN THE GREE..1'.i"HOUSE

FRO~I AMERICA.o'\l EUI SEEDS

srORED u-.lJER YARJOUS CONDITIONS

Storage

Pcr cent seedling production after

No. of years stated

Genni-

nadOR

Moistucc

prctfC"J.t- Te::mp., content

ment
C
pt:r cent

L:1b.

lAb.

momh
alScc.

-.

"

Scaled
Se:llcd

Scaled
Sealed
Sealcd

",6

Op~

-,

miner

,3
3

Nom:

Con-

'3
'7

"q
"
"

"

"

0
0
0

'4

"

'4

0
5

'3

,6

33

U)

~z

'5

"2Z

'0

'7

43

",8

Sealed

70

3
2

Scaled

55
58

'"82

75

78

8,

73

'5
5'

&>'1

Sealed

,3
-

",6

S,,"OO

33

,8

"

0
'5
'3
,8

"

....
2
'5

8,

83

55

3'

,8

Op=
Sealed
SCaled
S,,"OO

78
9'
87
80

58
70
57

5'
78

'5

"
67

73

,6

80

l.e:!l1: w:IS

Iillldc.

44

83
6,

6,

59
8]

68

7'

45

-=:3

75

AsHES

FraxiIJus(ash) seeds, also, 3rc among those which have been repor.h..d .to
lose viability quickly under ordinary storage conditions. G. P. Steinbauer
(I937), as a result of his experiments with FraximlJ, concluded that these
seeds are much more sensitive to moisture content than to temperatureat least, over a period of a year in storage. Taylor (I94I) reponed a serious

86

;3

0
0

Sealed

Indicatcs thaI no

'5

0
0

Smlcd
Sealed

5e-.llt:cl

"

~toragc for

LONGEVITY OF TREF,

SEEl)~

loss of viability of Fra:r:illlls seeds by the end of onc-:md-:l-half years of

storage at warehouse temperatures. However, Barton (19450) found that
seeds of Praxinus with moisture contents of approximately 7 to 10 per
cent may be kept for at least seven years by using scaled containers at Soc.

-=--

V.,rlLLOWS

Seeds of willows (Salix spp.) also lose their viability quickly when
exposed to the au. This has been assumed to be due to e.xcessive drying,
but Nakajima's work disprovcs this assumption. He (Nakajima, 1921)
found that seeds of Salix opaea, S.japonica, and S.l'dllii, retained their
viability much better in closed tubes uver a solution of 50 per cent by
volume of sulphuric acid in water than they did in the open air. In later
work he reported th:n seeds of Sali.T pi~Olii and S. japQlliCd in the open
air lost t.heir ability to germin.."lte within a \I eek, but when enclosed over
the sulphuric acid solution as mentioned above and stored at a low
temperature, ther still gave 53 per cent germination after 360 days of
storage (Nakajima, 1926). Such a solution gives a relative humidity of
only 13 per cent, which is much lower than the average humidity of the
aonosphere at the ripening time of the seeds. Evidently the injury in the
open air is not due to e.xcessive drying. The life of Salix caprea seeds, which
is normally onl)' thirty to forty days, can also be extended by drying the
sl."eds and storing them at a temperature of 6" to 9"C. (Janisevskii &
Pcrvuhina, 1941).
13IRCHF.S

Seeds of Betti/a letlla and B. poplI!lfolia- 'kept' perfectly during one

year of air-dry storage at room temperature, while those of B. papyri/era
suffered loss in viability (Joseph., 1929). The optimum moisture content
for seeds stored at room temperature in sealed containers lies considcra hly
below that of freshly haryesteJ seeds for B. papyriftra. (0.6 per cent), while
B. popu/iJolia seed keeps best with a medium amount of hygroscopic
moisture (5'2 per cent). B. lenta seed keeps well for one year in all except
very humid conditions. The moisture content proved to be unimponanr
during ~me )'ear's storage of the three species of Betula in a refrigerator.
POPL.\RS, ETC.

Engstrom (1948) recommends that cottonwood (Populus sp.) seeds be

planted immediately after harvest, since there is much Joss of viabilit)'
within two days. Seeds of American aspen (Populus tremuloides) and
balsam poplar (P. balsamifera), on the other hand, ga\'e vigorous
germination at the end of four weeks, and 45 per cent germination at the
end of eight weeks of storage in open dishes (~'loss, 1938). After eight
weeks, however, deterioration was rapid. Although these seeds have been

87

5Ef.l)

PRJ::SERVATIO'.... '\''-'1> LONGt:.\'lTY

reported nOt to tolerate desiccation) storing over C:llciurn chloride at - 5C.

showed remarkable extension of life, seed of the Russian poplar (P.
petr01J)Sk:yIJ1W.) giving 70 per cent germination after two years. Storage of
Populus tremuloides and P. grandidelitatiJ. seeds at room temperature in a
controlled-humidity series showed the optimum relative humidit}- to be
20 per cent, where seeds of the former remained viable for 555 and those
of the latter for 455 days, :lS compared ",ith a life-span of only 28 days in
the open air (L. P. V. Johnson, 1946). Busse (1935) presented evidence to
show that the loss of viability of aspen seeds is due to the injurious action
of oxygen; also that higher temperatures hasten the degeneration.

Norhofaglls mmziesii seeds stored in the open air will not remain
viable over winter, but in sealed containers at "20 co 50:C. full germination
was retained (Bibby & Williams, 1953)
FRUIT

TRl:ES

The ability of rosaeeou5 seeds to \,~thstand dry stonge is of great

practical interest, because, in the case ofspecies used for understock-such
as PrumlJ amaicaM ~1arsh-hea\'y crops sometimes appear only in
alternate years) which makes it desirable to carry over seeds for poorbearing years. Also, with rosaccous seeds that have a short after-ripening
period, it is convenient to keep the seeds in dry storage until the time for
stratification in mid- or late winter. In 1851, Fleischer stated that rosaceous seeds retain their \;ability in dry storage for considerable periods
(Fleischer, 1851). Seeds raken from apples and stored dry for two-and-ahalf years germi.nate almost as well and give nearly as high a seedling
production in soil as do seeds one-and-a-half years old of the same sorr
stored similarly (Crocker, t928; Crocker & Banon, 1931). However, both
of these show considerably lower germination pcrcemages than do seeds
only half-a-year old. Apple seeds that are to be used to produce understock, or hybrid-applc seeds, will keep well if ther arc removed from the
apples in the autumn, cleaned, dried, and stored until it is time to str.uif)
in late winter for: spring planting.
Seeds of the American 'Wild plum (Prullus a1lll'ric011tl Marsh) retain
more than 60 per cent of their power to produce seedlings after fifty-three
months of dry stonge at 7 to woe (Giersbach & Crocker, J932).
Optimum germination of apple, pear, apricot, cherry, plum, and sour
cherry seeds occurred after storage at a relative humidity of 50 to 55 per
cent (Solovjcva, 1950). Higher humidities were detrimental. Storing peaeh
seeds dry for one
decreased their value (Brase, 1948).
t appears
no long-term storage investigation under a "arie~' of
conditions has been attempted for fruit-tree seeds. Such an experiment

rear
tmt

88

LOl\'GEV1TY 01' TllEE SEWS

lfUuld undoubtedly reveal methods of c:\'tcnding the life-spull of these

Sttds.
A number of other tree seeds have been the subject oflongevity studies.
EUnIlyplUS miniata seeds collected in 1867 gave excellent germination in
1880 (tvIiiller, 1880). Successful storage of a number of Hawaiian forest
tree seeds depended upon lowering of the relative humidity and the
temperature (Akamine, I95I). These same conditions preserved the quality
:md the germination capacity of pecans (c. L. Smith et al., 1933; Wright,
I9flj Brison, 1945)
Q~JlNINE

The keeping-quality of CinchotuL seeds under various conditions is a

matter of importance in planting and breeding programmes of this genus.
It is necessar}' to keep a viable supply of seeds for continuous nursery
propagation in order to supply seedlings for field plantings of the trees.
Also, where e.xtensive hybridization is practised. seed storage methods,
which permit retention of germination capacity of the hybrid seeds until
the desirability of the trees produced from a sample planting can be
determined, are essential. This might mean a storage period of from four
to ten years. It has been reported that seeds older than 186 days at the
time of pl:lllting in Puerto Rico germinated poorly (Anon., 194oa). In
Uganda, A. S. Thomas (1946) secured 90 to 98 per cent germination from
fresh seeds, 70 to 80 per cent from seeds nine months old, 50 per cent
from seeds twenty months old, and only 5 per cent after the seeds had
been stored for thirty-two months. j\'Iorrison (1944) stated that Ci11chona
seeds lost about 5 per cent of their gennination power in about two years.
Storage was at room temperature in closed containers with calcium
chloride or sulphuric acid as a drying agent. No data were given. Air-tight
containers with calcium chloride also prolonged the viability of C.
Jedgerial1l1 Moens. in Puerto Rico (Kevorkian, (941). A refrigerator as
well as room temperature was used in the last instance, but little temperature effect was noted up to t,\clve months of storage, when 33'S and
30 per cent germinations were obtained, respectively.
In an article onCillcltOlld seed storage, Krcyer (1939) recorded information received from thc Biological-Agronomical Institure in Tanganyika to
the effect that the germination capacity of freshly harvested CitlcJlrJrza
seeds reached 99 per cent, Five months after gathering, however, the
viabilitj" had become reduced by" -l- pcr cent, and after eleven to fourteen
months only about 50 per cent of the seeds were capable of germination,
Krcycr stored seeds of C. [r;dgeriana for 418 days with various drying
agents in hermetically closed jars at a temperature of 25C. Seeds showed
a high germination capacity when kept in jars with largc doses of sodium
bromide, calcium chloride, a special desiccant called 'Adorosu" Or an
average dose of caustic lime. None of the controls germinated after 418
days.

-.
SEED PRESERVATION AI\'D LONGEVIn'

Cowgill (1944) has done extensive work on Cil/dlOna, part of whieh

dealt with seed storage for fifty-four weeks but with variable results.
Data are available to show the effects oftemperature, moisrurc cOntent,
light, and gaseous exchange,upon the keeping-quality of seeds of Cincho}/a
ledgeriana for periods up to and including four years (Barton, 1947)- Seeds
with 4,8,6'0, and 9-4 per cent of moisture were stored in open and sealed
containers in the laboratory, at Soc., and at approximately- ~4C.
Deterioration in the laboratory,,-as evident after eighteen months, and
germination was reduced to approximately one-half or less of the original
capacity after twenty-four months_ There was no advantage of scaled oyer
open storage at this temperature. 'Vhen the moisture content of the seeds
was as high as 9'4 per cent, there -mlS some evidence of harmful effects of
sealing and of light in the laboratory. Loss of germination capacity also
proceeded rapidl)' in open storage in a humid, sec. room. On the other
hand, seeds stored se."L1ed at 5eC., or open or sealed at approximately
-4C., kept perfectly during up to fony-eight months of storage. Within
the limits of this experiment, temperatllIe appeared to be a more important
factor than moisture content in the effects on Gi,lcllOna seeds.lt was concluded that C.. ledga"iana seeds can be kept succcssfullyfor at least four years
by maintaining a moisture contcnt of helow 9 per cent of the dry-weight
uf the seeds and keeping the temperature between SoC. and -4C.
(Barton 1947).
Tree seeds which will not tolerate drying ,,~11 be discusscd in the next
chapter.

--

. .:

.;;

go

,.

x
LONGEVITY OF SEEDS OF AQUATIC PLANTS, ETC.

IN a consideration of the storage in 'water of the seeds of aquatic plants,

t'.'.'o things must be kept in mind. First, only dormant seeds can be stored
in water at temperatures permitting germination, and second, many
seeds of water-plants of the temperate zooe after-ripen in water at temperatures near or several degrees above the freezing-point. When such
seeds are fully after-ripened, their germination may take place at the low
temperature, and will certainly occur upon transfer to a higher tcmpcra[me-provided, of course, that other conditions arc suitable and the seed
is viable. Long preservation of aquatic seeds in water] therefore, is
dependent on the dormancy of the seeds and persistence of a temperarure
\\'hich will not breaK the donnancy. The behaviour of Alisma plantagoaqWltica fruits illustrates both these points. If the intact fruits or seeds
arc placed in water and kept at room temperature, they will remain intact
and viable for months or even years on end with little or no germination.
If transferred in water to any temperature between 1 and IQ0e. for oneand-a-half months, they after-ripen and germinate freely when returned
to room temperature. They begin germinating even at the after-ripening
temperatures if left there for four months. These seeds al"o germinate
readily at room temperature if frozen and thawed repeatedly while full of
imbibed water, or if the seed-coats are broken mechanically. Freshly
harvested or dry-stored seeds behave in the same manner as those stored
at room temperature m water.
Isely (r944) nored the dormancy of fourteen species of Scirpus seeds
at maturil)', and the need for an after-ripening period during which the
seeds should be kept in a moist medium or in water at 2 to 4C. Many
species of Scirpus appeared to have an extended period of viability when
properly stored._
?vluenscher (1936, 1936a) stored seeds of thirt) genera and sixty-one
species of temperate-zone water-plants under four different conditions:
in water at 1 to 3e, in water at the laboratory temperarure, and in air
at these two temperatures. Germination tests were made in water in a
greenhouse after abont two, five, and seven months of storage in each
conditioIL Seeds of the following plants showed litde or no germination,
except after storage in water at 1 to 3e.: fourteen species of Po/amogetoll, three species of Najas, Alisma plalltago-aquatica, three species of
Sagittaria, Butomus umbel/atus, Zizania aquatica, nvo species of Scirpus,

9'

Sf.D) PRESERVATION ....ND WNGEVITY

Acoms calamus, Calla palm/fis, Qronti1lnl aquaticum, Pcltandra virgil/ien,

Heteramhera dubin, Pontederia cordata, Erio[Qulon uptangulare, Polygonum amphibium. Ceratoplryllu11I demersum, Nympho::.anthus ;;ariegatus,
Nymphaea tuiJerosa, Trapa. no/am, Elatine americana. and Lobelia dartmanna. The percentage of germination generally increased with rhe
period of cold storage from two to five and finall)' to seven months. No
doubt some or all of these seeds were dormant and were after-ripened by
storage in water at I" to 3"C. This would account for the increased
germination after the longer periods in cold w:lter. It seems that even
seven months of storage was not sufficient to produce full after-ripening
in some of the seeds.
Seeds of the following species showed little or no germination after
any of the conditions of Stonge used by Muenscher: five species of
Sporganium, Najas minor, Sagitlc1Tia latifolio. Lopkotocarpus spungiosus,
three species of Scirpw, and /(0[1Ior;s coh'a. Wher:h.er this was due '1"0
poor seed quality or a failure to provide proper germination conditions is
not evident from the data.
Muenseher found that Glyceria striata and Nasturtium ttJUlurliumdquot;rum showed fair [Q good germination after ill four conditions of
storage. Vallisneria americana genninated readily in water at the lahor:1tory temperature, whether it had previously been slored in cold water or
not. Of the spC1:ies tried, only one germinated after dry storage.
Guppy (1897) gives the records of the behaviour of the seeds of a
number of aquatic plants when stored in air and in wateI:. Set:ds of
NuplwT luleum and Nymphaea alhaare soon killed by drying. Seeds of the
former germinate more readily than those of the latter, but some arc
delayed until the second rear or later. Potamogetol/ seeds are also injured
by protracted drying. Howe\-cr, seeds of the following water-plants
endure drying without injury for the period mentioned after each, and
this is the longest period determined by Guppy unless othenvise noted:
Myriophyllum spicatum, I ! years; M. alt"Tllijlorum, I year; Lz'mnattthemum
lI,YmpIUlCQides, 2! years; S parganium ramostlJII. S. simplex, and Zamliclullia
palustris, 41 months; Callitriche aqllatua, 2 years, but rolted after drying
years; and Rammculus alfUlllilis and R. hedrroCl!us, many months.
DI'}ing for two months promoted the germination of Crrotophyllum
dcmersum, and drying for three or four months increased the germination
of Sagittaria sagittifolia seeds.
Seeds of Alisma plantago-aquatica, Typha /lJtifolia. and BUIOttlU!
umbellatus, stored dry in em'elopes in the laboratory at Yonkers, New
York, were tested for germination by breaking the fruit and seed-eoats
and placing in water at a daib' alternating temperature of 15 to 30C.
Alisma r2-!, 6~. 3t, and 21 years old gave 0, 40, 39, and 51 per cent
germination, respectively; l)pha 12!, 5~, and 4! years old gave 0, 96,
and 78 per cent germination; and Butomus 7! and 3! years old gave 9 and
26 per cent germination.

3t

9'

LONGE\'ITY OF SEEDS OF AQUATIC PU"l'-'TS, ETC

It has been shown by Haigh (1940) that seeds of the water hyacinth
(EicMwmia crassipes Solms.) remain viable in water in the laboratory for
at least five ye:l.fS, but that dry seeds f.'til to germinate after three years of
storage. Seeds of this plant were able to survive one month in ice at
approximately -4C., and as long as two months at temperatures up to
15C. (Barton & Hotchkiss, 1951). At 40C., also, the seeds not only
survived for two months, but gave excellent germination after such
storage. "When the storage period was lengthened to seventeen months,
however, 20 and 30"C. proved better than 5" or 4oC. for keeping the
seeds v.iJ.ble. The life-span of the seeds under these various conditions
was not determined, but the requirement of a combination of high temperature and light for complete germill:J.t.ion was established. Hitchcock
et al. (1949, 1950) kept water hyacinth seeds in water in the laboratory
for more than two years without germination. A knowledge of the
germination behaviour and the life-span of seeds of Eichhornia is important in the control of this plant in waterways, as enough seedlings may
become established in a year to pose a serious threat of reinfestation
following successful control by chemicals of the vegetative parts of the
plants.
Longevity is also an important factor in a control programme for
Halogeton glomrratlls, the seeds of which decline in viability after one
year (.>\.non., 1953).
Certain species of Polygomlfll can live either in water or on land, and
are imporrant as food and cover for water-fowl and fish, bur., under same
conditions, these same plants may become tiresome weeds. Viability and
dormancy in three Po/ygoflum species, P. amplzibium L., P. coccineum
Muhl. and P. Ilydropiperoides Mich..x., have been studied by Justice
(1944) Achenes of all three species placed under outdoor conditions
similar to their habitats lost their germination capacity during the first
winter. Also, seeds of all three were killed by a year of dry storage in the
laboratory. They could be stored successfully for longer periods in water
at zoe, where any dormancy present was broken.
Duvel (1905) found that wild rice (ZiZl17lia aquatica) seeds lost their
germination capacity if the)' were allowed to dry in the air for even a few
days, but that they retained their viability perfectly until spring if stored
in water at 0" to IDe. In the spring they must be transferred from the
storage water to the water in which they are to grow, without being
allowed to dry. Seeds of ,,"-ild rice are dormant when mature, and storage
in water near the freezing-point will after-ripen them while maintaining
their viability.
The quality of cultivated rice (Or..Yza sativa L.) grain stored under a
variety of conditions was reported on by Kondo and his co-workers in
a series of twenty articles published over the period 1927-38. The proper
amount of drying is essential to the maintenance of satisfactory foodqualities as well as viability. Drying by the use of C<l.lcium chloride (Kondo
0

93

SF.[[) PRESERV.\TION A.''D LONGEVITY

& Okamura, 1931, 19341', 193, 1935; !\!lcFarlane el aI., 1955) or calcium
oxide (Kondo & lsshiki, 1936; Kondo & TCras::l.ka, 193n) is recommended,
though heated air can also be used (Kondo & Ol-amut:l, '932-33, 1932-

330; Kondo el al., 1938). A more modern method is pre-harrest drying

with chemicals sprayed from an airplane (Anon. 1953a). The safe moisture content of rice grain depends upon the storage temperature (Kondo
& Okamura, 1938). Rice seeds stored for five years in a concrete silo,
hermetically seale<! (Kondo & Okamura, 1937), or for t..y cnty-tbree years
on a mountain (Kondo & Okamura, I93), retained satisfactory culinaI)
qualities, but were not capable of germination. Unhulled rice stored for
about 100 years in a granary was still edible, though it had an unpleasant
taste (Kondo & Okamura, 193&).
Sampierro (1931) was able to keep rice sds fully viable (99 per cent
germination) for eight years provided they were dried to a moisrure
content of 5 per cent and sEared in aD atmosphere of nitrogen. In atmospheres of carbon dioride or air, or in a partial vacuum, the seeds were
killed. Thirteen per cent moisrure was faw under all conditions. Similar
results were obtained by Saran (1945). Dore (1955) stored rice grain in
an air--eundirioned room of low temperature :md humidity, where the
seeds retained their ..;abiliry for three-and-a-half years. They remained
viable for two years in sealed containers with C:llcium chloride.
We have seen that seeds of some water plants m:n not survi~e storage
in the air, while others will stand severaJ years of dry storage in the air
without injury, and Nelumbo (see Chapter I) seeds live for some cenruries
in dry storage.
SEIDS L"JtJRED BY DESlcc.UIO:-.l

Some seeds lose their germination capacity in a very shott time when
tbeyare kept in the open air after harvest. This has been assumed to be
due to the drying effect of the air. As we have seen in Chaptet IX, manf
seeds, such as those for example of the American elm and some aquatic
planrs, which are supposed to be killed by drying, tolerate dcsiCC:ttion to
a remarkable degree and can be kept for long periods in a dry state at low
temperatures. However, there remain certain seeds which appear to be
killed by drying. Though mOSt of these are of tropical or semi-tropical
origin, there are some temperate-zone forms among them.
According to H. A. Jones (1920), the seeds of the river maple (Acer
saccltarinum) arc killed by relatively slight drying. \hen they fall from
the tree in June, they bear about 58 per cent of water. Regardless of the
temperature of exposure (0'" to 35"C.), they were killed when the
moisture content reached 30 to 34 per cent. 111 Jones's experiments it
required six days at 35"C. and ninety-two days at ODe. to reach this water
content or the death point. When these seeds were stored in a closed
vessel over water at the freezing-point and provision W:1S made for preventing carbon dioxide accumulation, they retained full viability for r02

9+

LONGEVITY OF SEEDS OF AQUATIC PL.4.t'\;1S, ETC.

--

days, which was the limit of the test. The low temperature prevented
germination and reduced the ratc of metabolism. The latter is an important consideration in any seed with high moisture content. River
maple seeds should be 5m'.l1 immediately after harvest. If this is impossible, because of the necessity of shipping or for any other reason, they
should be kept ncar the freezing-point, and water loss prevented.
They are not dormant hut begin germination in nature as soon
as they reach the moist ground. The seeds of the fall-fmiting sugar
maple (A. saccharum) show very different behaviour. They endure
complete air-drying and respond to about three months' low-temperature
stratification for eliminating dormancy.
Acorns (the seeds of oaks, Quercus spp.) have a very shan life-span
under ordinary conditions of storage (Oppermann, 1913; Gardner, 1937),
but extension of gelmination capacity up to ten months results from
mixing with dry sand and storing in ail-tight cans at 32 to 40F. (Mirov,
1943) Seeds of the tulip-tree (Liriodclldroll tulipiftra), which are injured
by drying, have been kept for four years without loss of viability by
storing in soil (Paton, 19H). Six thousand pounds of seed were placed in
a pit in two-inch layers alternating with layers of sand to cover the seeds.
A concavity to catch water was provided at the top of the pit, so that the
seeds were kept moist.
Kidd (! 914) attributed prolongation of the life of rapidly deteriorating
Hroea brasilimsis seeds to the presence of 40 per cent of carbon dioxide
produced by the respiration of the seed!> in a closed flask. He did not make
moisture determinations of these seeds but stated that they had a high
water content. He summarized, in pan, that 'the resting stage of the
moist seed is primarily a phase of ll:ll'cosis induced by the action of carbon
dioxide' (Kidd, 1914, p. 624). This is contrary to the claim made by H. A.
Jones (1920) that Acer sauhari1lum seeds kept bener when carbon
dioxide was not allowed to accumulate than when it was.
Busse (1935) thought that the rapid loss of viability of poplar seeds
when left in air was due to the injurious action of oxygen. He was able to
demonstrate that seeds in sealed containers with reduced oxygen pressure
remained viable longer than those subjected to full atmospheric pressure.
Sugar-cane seeds degenerate rapidly when stored in the open air. This
makes it impossible to ship them with assU1':lIlce from one sugar region
of the world to distant ones where seedlings arc desired for breeding.
Verret (1928) found that \~abilit}' eould be lengthened materially by
mk.ing the seeds from the thoroughI}T air-dried heads, placing them in
cans with 9 gm. of calcium chloride to ! litre of space, displacing the air
with carbon dioxide, hermetically sealing, and storing at the freezingpoint. In these seeds, low and perhaps constant water-eontcnt and absence
'Str:nifiCition' is a term derhed from the pr:u:t:ice of nurserymen (0 Rpread
layers of seeds alternating: with byers of moist soil or sand for o,er-winter or other
low-lempennure treatment of seeds. The same term is now generally used to denote
any low-temperature lre:Itment of seeds in a moist medium.

95

SEED PRF.S.RVATION ."'-.'\TI LONGEVITY

of oxygen st;:em to be necessary for retention of germination capacity. It

is possible :lIso that C:lrbon dioxide may playa positiYe role rather than
having its ,'aluc in merely displacing o:\'-ygcn. These results were confirmed
in the main b) Darragh (1931). More re<:ently, freezing at o"F. has been
added to drying for successful Storage of sugar-cane seeds (Abbott, 1950;
Anon., 19518, 1952).
Citrus seeds are among those which arc injured by df!iog and hence
deteriorate rapidly under ordinary conditions of storage. Dealers in these
seeds have experienced difficulty in maintaining their germinatiye power,
although Florida sour orange seeds have been reported to be more
resistant to drying effects than most other citrus seeds. For Ille most part
the seeds have OC'eD kept moist, either in the original fruit juice or in
moist sand or some other moist medium after cleaning, until they were
sold. Inquiries from scedsmen, together with the bck of experimental
wo:'"k on the keeping quality of these seeds, led to a study reported by
Bar[Qn (I9.HQ). Seeds of grapefruit, sweet orange, sour orange, and rough
lemon, were s[Qred under different conditions to deh..Tmine factors which
would prolong life.
Work with grapefruit seeds has shown the importance of moisture
content, temperature, and scaling effects, to viability. Hi~h moisrures
(roo to I31 per cent calculated on the basis of dty-weights of the seeds)
were detrinlental to seeds in c100ed containers, especiall)' during stOrage
at laboratory tcmper:lIures. However, drying of grapefruit seeds to 60 per
cent of the dry weight, which is still a ....ery high moisture content, prolonged their viabili~' in scaled Storage in the laboratory as comp:rred with
open storage at this temperature. Ob'\;ousiy, then, SOffie factor other dun
carbon dio:Jde accumulation was operating to prolong the life of grapefruit
seeds, as sealed storage W:lS not favourable in all cases. This was shown
further b)' the less fa\'Ourable effect of sealed storage at SoC as compared
with open storage at that temperature. Mould appeared at an early d:ue on
moist seeds in sealed storage in the laboratory and may have been aiactor
in deterioration, but it wouldbe \'ery difficult to say whether the mould led
to deterioration or whether deterioration led to the development of mould.
The best storage condition found for grapefruit seeds ""as in open
containers in a room held at 5C. After a year of open storage in this
very humid SoC room, grapefruit seeds contained 18 per cent of
moisture. As limited tests indiC:lted that fresh seeds wcrc somewhat
injured hy drying on blotters in the laboratory until they contained S2 per.
cent of moisture, the beneficial effect of drying to I8 per cent at Soc. is
of particular interest. Ccnainl)' it cannot be said that their longevity
depends entirely upon water coment. It appears that the drying temperature and perhaps oilier factors such a.c; rapidity of desiccation, etc., arc of
importance. In this respect, these seeds differ from those of river m:lple
a.c; described by H. A.. Jones (1920). GtrUS seeds apparently ,,'ary Vriddy
in their tolerance to desiCCltion. "llercas the germination of seeds of

96

LONGEVITY 01' Sf.f.DS Of: AQU.-\'rIC 'U.NTs., EtC.

grapefruit \\-as reduced 30 per cent by drying to S2 per cent of moisture,

and that of sweet orange seeds was reduced 80 per cent by drying to 25
per cent of moisture, sour orange and rough lemon sceds could he dried
to 4 per cent of Illoisture and still retain 25 and 50 per cent of their original
germination capacit)t.
.
The harmful effect of a temperature of approximately ~ Soc. on the
keeping quality of seeds of grapefruit, sour orange, and rough lemon, is in
direct contrast to the efficacy of this same temperature for maintaining
the viability of many seeds which e:lll be dried to relatively low moisture
contents. That the deleterious effect OIl citrus seeds of such temperatures
was not due entirely to high moisture content was ucmonstrated in the
experiments of Barton (1943), which showed that rough lemon and sour
orange seeds dried to 4 per cent of moisture and still capable of germination, were unfavourably affected by below-freezing temperature. Childs
& Hrnciar (1949) believed that micro--organisms play an important role
in the deterioration of citrus seeds in storage. They stored large lots of
thirty-four varieties for si.x months and of thirty-three varieties for eight
months at 3SoP. (I7e.) in moist sawdust or moist moss in unsealed
containers, after dipping the seeds in a 1 per cent solution of 8-bydroxyquinoline sulphate. Twenty-nine of the lots gave better than 90 per cent
germination after six: months and the average germination after eight
months of storage was 845 per cent. There were, however, great differences between varieties. Deterioration of citrus seeds in storage was
attributed to micro-organisms, moisture, temperature, and aeration.
Richards (195z) found a marked reduction in viability of seeds of Country
lime and Bibile sweet orange when they were stored at normal air temperatures of 78 to SoF. for more than three days after extraction from
the fruit. The interrelation of gaseous exchange, moisture content, temperature, and micro--organisms, in their effects on the keeping quality of
short-lived cirrus seeds, is not apparent at the present time. Much more
work is needed to clarify the situation.
The rapid deterioration of seeds of maga (Molllczuma- spet:iosissimfl.
Moe. & Sesse) after harvest, coupled with the economic importance of the
tree in yielding wood of good quality, has made it desirable to determine
conditions for keeping seeds viable over longer periods of time than they
usually live. Germination occurs shortly after the seeds fall from the tree,
and may even start before the fall of the capsule. From this characteristic
one would judge that desiccation might destroy the life of the embryo,
which is large and occupies aU of the space within the seed-coat. Seed
storage of maga was the subject of a study made by Barton (1945). It was
found that the life-span became reduced significantly within two weeks
under ordinary storage conditions. "'hen the moisture content was as
high as 33 per cent of the wet weight, the seeds could be kept for a month
in scaled storage at Soc. Drying at the laboratory temperature to
approximately 10 per cent moisrnre reduced the germination capacity
G

97

SEED PRESERVAtiON .\>.''D WNGEVITI"

immediately by one-hair, but perroined subsequent retention uf this

lo.... ercd germination capacity at - 5". 5. or lOe. for at 1~1: six:
months, and possibly longer.
These results are in general agreement with those of Marrero (1942),
who dried seed-lots of maga with the aid of an electric fan to 87'5, 75.0,
and 62'5 per eent of the origin.a.l weight.
Refrigcration is necessary to extend the life of many tropical h:lrl.lwood
seeds from two or three months to one year (Marrero, 1943). Seeds of the
Cupuliferae are lurmcd by dry storage, but mar be kept for as long as nm
years if lhey are mixed \\-ith river sand and held at 0 to 5"C. (Kondo ,t al.,
Ig.p). The viability of air-dry seeds of rnangosteen (Careillia t/1angostana
L.) is lost after four weeks' storage, but may be preserved for seycn to
eight weeks in moistened charcoal or peat at room temperature (Winters
& Rodriguez-Colon, 1953). Storage for onc weeL: O>er calcium chloride
killed all the seeds. Other tropical seeds must be kept moist to maintain
viability, and thus do not have long life-spans in storage. Among these
are seeds of cacao (E\-ans, 1950), coffee (Gonz:oilcz-Rios, 1929; lJltec,
1933; Anon., 19410), ErytbroAy/olI and Litchi (McClelland, 1944), brgeleaf mahogany (Lopez, 1938), and Macadamia (Chu et 0/., 1953).
D. G. W'hite and his co--workers have investigated the storage requiremcnts of bamboo sceds (D. G. White. 19+7; D. G. White & Villafane,
1947; D. G. White & Delgado, 194-8) and have found that the most
practical method of prcsen-ing their viability is storage over calcium
chloride at room temperamre, though rcfrigcration results in still greater
life--extension. Webster (1g..+8) tried several methods of storing seeds of me
tung (A/curitrs) trec; but the only one v. hich prevented loss of viability
was stratification with moist sand followed by storage in boxes in a cool
building. Under these renditions full germination capacity was rctained
for sLx-and-a-half months. The life of these seeds is greatly prolonged by
storage at 3+ to 40F. according to Large ~t a/. (1947), but Shear & Crane
(1943) reponed that mng sceds stored at 7C. lost their viability soonCf
than did seeds from storage temperarores which ranged either from 23
to 32C. or from 7 to 55C.
It has been sho'W"Il abm"c that, although seeds of all kinds deteriorate
very rapidly in the tropics, it is possible, through humidity and tcmperanue control, to extcnd the lif::-span of many valuable seeds, thns permitting the correlation of direct seeding with correct planting schedules.

-.

..".".-.:."
.0.
-~
: ,',; - _ .

~-

Xl

EFFECT OF PACKETS ON SEED VIABILITY

IT has been well established that moisture content and temperature
determine, to a large extent, the keeping quality of seeds in storage.
Neither moisture nor temperature '-"all be considered alone, as the effect
of onc depends upon the other. These factors not only affect seeds while

they arc in controlled storage rooms maintained by sccdsmcn, hut

continue to exert a vital influence after secus are removed and packeted
for the trade. Furthermore, it is essential that the seeds maintain full
viability under the original storage conditions, in order to withstand
further storage under unfavourable conditions. The better the stOf:lgC
condition for keeping the seeds viable, the better the chance of sUIvi\'al
when they are removed to an unfavourable environment. Other conditions
being equal, low temperature has been found to be superior to higher
temperatures for the maintenance of germination capacity. Howeycr,
seeds dried and stored in sealed containers at ordinary room temperature
may kcep better than those stored open in a room at Soc. where high
humidity prevails, as has been demonstrated repeatedly in storage
experiments at Yonkers, New York. In these cases, tilen, resistance to
further storage .....'ould be in favour of the original room-temperature
storage or the storage conditions permitting the highest retention of
germination capacity.
The keeping quality of any particular lot of seeds does not depend
upon its initial high quality, bur upon the storage conditions. Seeds of low
germination capacity may be stored successfully for fairly long periods if
the storage is appropriate. On the other hand, seeds in which deLCrioration
has been initiated, even if the germination capacity is still high, arc
incapable of remaining viable for long periods under adverse storage
conditions.
All of these facts point to the necessity of maintaining rooms in which
temperature and humidity can be controlled, if commercially important
seeds arc to be held over for sale in subsequent years. Responsible seedsmen have come to realize this and have provided the proper facilities.
Howeyer, another problem, that of deterioration upon removal from
storage, is being recognized as of the utmost significance. Seedsmen must
packet their seeds for the trade well in advance of the time the consumer
buys and plants them. Kraft paper envelopes are commonly used, and are
sent to wholesale and retail dealers in many parts of the world. Some of

99

these packeted seeds are stoted in warehouses or retail stores under

conditions of high temperature or humidity, or both, l here :;erminarion
capacity dccte:lSCS at such :l rapid rate that the seeds may be worthless
when the customer finally plants them. The rapid deterioration under
unfavounble conditions of onion seed upon removal from storage was
reported by Barton (1939, H}p).
Some idea of the importance of the sced-pacJ.:et industry and of the
poor quality of seeds found in manysueh packets may be gained from aseries
ofarticles by M. T. MUDD and his co-workers, published between 192., and
19+9. Two of these '\\i.ll serve as ex:lmples (Munn et 01., 1929; Munn &
Munn, 1936). In 1926, 1927, and 1928, packct<; of vegetable seeds were
purchased from forty-seven different seedsmen or dealers in New York..
Out of C)6., packets, 9'2 per cent were worthless for planting purposes. In
addition to these, there was a relatively high percentage of packets of very
litde seeding value. In 1923. 23 per cent of 347 packets. and in 192." 18
per cent of 600 packets, were worthless. This represented very serious
losses to the growers. The analysis, by these workers, of the germination
behaviour of the sccds~by firms, with an aetuallisting of the firms and
the quality of seeds they sold~has doubdess contributed to the forcing of
some unscrupulous dealers out of the I:IW"kct.
An analogous situation has been found for flower seeds in New York
(Munn & MUlUl, 1936). In 1936, of 500 packets of flower seeds examined,
57 per cent were excellent, good. or entirely satisfactory, ",hile the
remaining 43 per cent were unsatisfactory. Fifteen per cent were worthless.
Similar work has been reported from the .Massachusetts Agricultural
Experiment Station (Snyder & Tuttle, 1937), from the Seed Laboratory
of the Department of Agriculture, Canada (Dymond, 19(7), and the
California State Department of Agriculture (Suslcin, 1953).
As a result of these and probably many other unreported tests, there
arc now Federal and State seed laws in the United States which seek to
govern the type of seed which is offered for sale. From the fundameutal
seed behaviour described in previous chapters, however, it is e...i dent
that the problem of control is dependent upon a basic knowledge of the
response of seeds to stonge conditions-not only in the special storage
room of the responsible seedsman, but also from the time they lca.e that
room until they reach the grower, and, ultimately, the soil.
Among the published reports on the germination of packetcd seeds is
that of E. Brown & Goss (1912). (r,-cr a period of five years (from 1907 to
(911) they tested 18,571 packets of.egetable seeds. Thesc werc the familiar
paper packets lithographed in colours and put up by sixty different firms.
These paper packets ate shipped in boxes and hence arc known as box
seeds. Mail-order seeds from nt"enty firms were also included in the stUdy.
A marked difference in the quality of the box and mail-order seeds was
noted. Whereas the average germination of all of the former was 60'5 per
cent. that of the latter W:l$ 17'5 per cent. When the germination of box
'00

EFFECf 01' PACKETS ON SEED \'IAnn .ITY

seeds by firms was analysed, the difference in average gcrmin:J.tion for the

different firms was marked (from 8T"S to 36.5 percent), while the difference
in average germination of seeds from mail-order houses was small (from
Sr8 to 70'3 per cent). Differences in box and mail-order seeds of the same
variety ranged from 2"3 per cent for parsnip and 3"3 per cent for spinach,

to 28S per cent for cabbage and 29"5 per cent for onion. The average

-=

--

germination of box seeds put out by anyone firm was uniform from year
to year, but the differences in qualiryvaried from 73"7 per cent for one firm
to 42-8 per cent for another. Many instances were found in which there
were statements of high germination on the packet 31ld yet the authors
obtained only 42-8 to 43.6 per cent germination. They concluded (E.
Brown & Goss, 19I2, p. 9): 'It would thus appear that such firms either
do not tcst their seeds accurately or else disregard the results of such tests
to the extent of including lots showing unsatisfactory germination in tbeir
output of packeted seeds for the box trade.' The dependence of mailorder firms on satisfied customers was given by Brown & Goss as the
reason for the superiority of theit seeds.
In the light of the evidence nO'w available, the better qualit}' of the
mail-order seeds might be attributed, at least in pan, to the direct route
from the fayourable storage rooms of the seedsmen right to the consumer.
Such seeds would tend to be subjected to possible high temperature and
humidity conditions for a minimum period and hence haye a higher
germination capacity at planting time than the box seeds. Also, it may
very well be that secds which may have given 80 to 100 per cent gcnnination at the time they were packeted for distribution have lost their value
entirely before they reached the grower, if the wholesale or retail establishment has poor storage facilities. While this fact may remove some of the
blame for seed failure from the secdsm:m, it is still his responsibilit}, to
proyide the best possible seeds for filling the packets he sends out.
As the moisrure content of the secds is such an important factor in their
deterioration, and as the paper packets commonly used in the trade are
not in any sense waterproof, it became of interest to determine the
difference in viability of seeds in these packets as compared with ones in
waterproof packets. The results to be presented below have demonstrated
the superiority ofcin cans with tight-fitting lids, or ofheat-sealed aluminium
foil envelopes, over the ordinary kraft paper seed-envelopc for the successful racketing of onion seeds (Barton, 1949).
Germination tests and moisture determimtions were made immediately
after receipt of seeds of onion (Allium (epa L. var. Ebenezer). The
moisture content of two portions of the seeds was adjusted to Io6 and 35
per cent of the dry-weight of the seeds by placing the seeds in desiccators
over water or calcium oxide until the desired weight was obtained. The
seeds were then stored at 5" and 2S"C. in open and in sealed containers.
ViabiJity tests were made after 3, 6, 12, and 24 months of storage. At
the end of each of these periods, seeds from each storage condition were

ror

SI'.ED I'RKSERVATIO:-i A.I'\'D LONGEVITY

racketed in papcr seed-envelopes of the type llsuaJIy employcd in the

trade, and iu tin cans \\ith tight-fitting: lids scaled with sealing Will:.
These packets and tin cans were then placed for further storage in a
0D
room at 3 C. The paper packets were stored in a desiccator over a
saturated solution of sodium chloride giving approximately 76 pcr cent
relative humidity. Fi"e packets and five tin cans of seeds from each
storage condition were prepared at the end of each of the four storage
periods, to allow for resting after from two weeks to six months at 30DC.
In a few cases, seeds from packets and tin l:-'1lllS 'were also tested after nine
and twelve months.

At each testing period, duplicates of 100 seeds each were germinated

(i) on moist filter-paper in Perri dishes at 2o"C., and (ii) in the greenhouse
in a mixture of equal parts of &1.nd, sod soil, and b'T.lnulared peat-moss.
In Table XIII are shown the germination percentages obtained, both
at 20"C. and in the h'1'ecnhouse, from seeds packeted after original storage
periods of 3, 6, 12, and 24 months at 5" and 25"C., and then held for
additional periods of 2 weeks to 6 months at 30 De. and 76 per cem
relative humidity.

TABLE XIT!
GER.I\HNATION 01' STORED ONION SEEDS AI'TER PACKETING AND FURTI-/f.R
STORAGE

Gc:nni_
nation
condition

Time

;"

Teml' Open

in <

mouths

"

sc:alcd

J\1ois-

,
's

Mois!

filter_
[laper'll

zo~c.

'3

"

5
5
0
S
S

106

,s

106
3'5

0
S
S

10'ti

0
S
5

106

0
5
S

35

3"5
106
35

0
S

5
S

Gcrmination~

<=

'S

106

3'5

% Gennin:llilm afwr fllrrh~T stQt:I;;:e in

Original ~tQt:lge condition

,S

packets at 30~C, :;6% relalive:

humidity
Paper packets
Tin cans
(months)
(months)

,
""

6,
4'
6, ,6

97
96

"

6,

97
97

"
" "
97

<

"

6,

94 93

77

,
,

,
" ,
H
" ,
33 ,
3 ,
,
45

97

9' S,

94
93
9'

93 9'
9 6 94

9'

94 9J 7' 53
86 79 6
9' 94 B,
'4

93
~,

95

96

93

"

J8
45

95

"

98 93 78 37

" ""
94

lD2

,
,

S' "

3 6

q; 9'
94 9'

"96
94
"
')6 ,B
93 94
93 94

97 9'

, "

,6
86 53 '5

95 77

, ,

0'5

97
97 97

,S

97
97
97

93

97

95

9'

93

97

93 97
93 95
97

"

9.1
94 q6
9'
97

"

,6 93 9' ,~
94 75 77 8,
96
9' 94 9J
94

'"

" 9'
"" :J "

97
97
97 99

96 94 9'
95

94

--

EI'l'Ecr OP PACKETS ON SEEU VIAIJILI1Y

=
=

"

1}2

106

,6

3-'

91

10'U

3"5

S
5

o
o
s

s
3

Soil in
m-ccn-

huu..e

,
,

::

8<)

93

1)0

84

1)0

<)

62

7'

(H

53

59

90 90 89 8.. 94

820 32

1)6

8S

9.3

95
97

95 95 26
94 ')0 29

9 2 97 96 93 92
92 l}l. 97 94 96
95 1)6 95 <)2 93

88

33

<)

83

45

20

38

10

79

<)

84
106

71

3'5

68

I}l

87

14

84

17

29 12 0
8 <.>
28170

85 79
81 64

2.

85778 1

50

8,

87 83

84

71

o
o

8,

""

87 8.;

8,

93

70

61

25

89 78
76 ;!~

84 84

\IV

78 86 7'

JI

5
S

84 85

73

83 85

67

93 113
Sl 8,;:
8.j. 8-1-

77 80
8, OS
ii U

,. 88

87 80

93

81

7'1.

85

77 75
"" 8,

88 84 84 78 81

61

62

'}O

')0

S
5

8,

77827864253

92
77

8,;: 85 62 ~_3
86 80 76 -6

,
9

86
44-

81

<)

86 81

86 8,

77

70000

40 47

37

17

75

"

24

,6

5
6

87 S8

10,6

3"5

,.

3620

"7
2
8857

68

12

28

818163910

8,

o
o

o
o

59
liS

,.

28

8) 86 86 87 89

.",.

106

"'
rcmO\'<I1 from original storage condition for pad:cting.

.. Per cent germination UpllD

- Indicates WI no test was m:J.dc.

It will be seen that almost complete capacity to germinate on moist

filler-paper at 20"C. was maintained under all the conditions of the

original storage up to and including twelve monLhs. This ffit..":ln5 that for
storage periods of one year or less, temperatures of 5 and 25C., and
open and sealed containers, were equally effective in maintaining a high
quality in the onion seeds as measured by their abili~' to germinate
immediately upon removal from storage. Furthermore, for germination
under these conditions, there appeared to be no advantage in the initial
drying: of the seeds from 106 to 3-5 per cent moisture.
An extension of the sturage period to twenty-four months, however,
proved deleterious to the onion seeds containing 10-6 per cent of moisture
when the storage temperature was 25C. This will be seen in the reductioll
of the germination at 20C. to 76 per cent. Widl this one exception, the
germination capacity of all of the seed-lots immediately upon removal
from the original storage conditions was high, ranging from 91 to 99 per

'"3

52
72

87 1)0 85 85 79
84 S8 90 16 ,6
90 91 88 78 82

3"5

52
8,;:

SEED PRESERVATION M'D l.ONGI::vrn

cent. That thcre were differences in these seeds as regards their abili~' to
withstand funher unfavourable storage \\;!l be shown below_
The advantage of open storJgc at 25"C. over sealed storage of seeds
containing 10.6 per cent moisture is evident_ The atmosphere in the 2S"C.
storage chamber was dry enough to permit a moismre content 10"er than
10-6 per cent for the seeds in open cont:1iners. It always has a worse cffect
on seeds to seal them with e.xcess moisture than to leave them in open
containers in atmospheres of low humidity.
Reference to the data secured by testing these same seeds afrer further
storage at 30e. and 76 per cent relative humidity, reveals the striking
superiority of sealed tin cans over the regulation seed-packets (Table
Xl II). Six months after packeting in tin cans, a \-ery high capacity to
germinate on moist filter-paper at 20C. was rerained by all lots, regardless
of the original storage conditions or time. e.xcept by the one with reduced
germination at the time of packeting, and e\-en that lot still gave 59 per
cent germination. An extension of the time in tin cans at 30C. to t""'elve
months failed to lower appreciably the germination capacity of onion
seeds which had received a pre,,;ous storage of six months at SO or 2SC.
This is in striking oontrast to the filte of the seeds in paper packets, all
of which were entirely worthless in si.x months, and were already so
reduced in germ.in::Ltion \;gour after three months that good seedling
stands could no longer be obtained in soil.
Most rapid degeneration in paper seed-packers is seen for the seeds
containing an original 106 per cent of moisture and from an original
storage temperature of 2SC. in sealed containers, regardless of whether
the original storage time was for 3. 6, 12, or :q. months. This was to be
expected as these were the most unfavourable original storage condirions
used (Table XlII). These seeds germinated 96, 91, and 93 per cent
respeeti"'ely after 3, 6, and J 2 months original storngc, and wcre indistinguishable in this regard from the seeds stored under all of the other
conditions. The inferior quality showed up. however, in the decreased
resistance of the seeds to funher unfavourable conditions-in this case
paper packets at 30e. and 76 per cent relative humidity. This response
was similar to that obtained previously (Barton. 1939. 194.1).
Reducing the moisture content of onion seeds from 106 to 3-S per
cent before storing in sealed containers at 25C., made possiblc the
preservation of their germination \;gour for somewhat longer periods
upon removal to 30C. and 76 per cent relative humidity_ Some of these
effects, together \\;th a comparison of germination capacity in soil and on
moist filter-paper, and of seed-cm'e1opes and tin cans for packeting, arc
shown in Fig. II.
SC'o'eral facts are e,,;dent. One of these is the increased tolerance to
subsequent unfavourable storage of seeds stored with a low moisture
conteur; another is the efficacy of tin cans for satisfactory keeping after
packeting; and still another is the poor performance of weak seeds in the
14

EFFECT OF PACKETS ON SUD VIABILITY

soil as compared to moist fi1cer-paper. Again, it should be emphasized that

only those seeds which had had previous drr storage, :lud which had

'co
'0

w
'0

"
'0
'0
'0

Tin Cons

z
~

'0
'0

'00

3.5

10.6

w '0

~
w '0

Poper

Seed Pockets

'0

"
>0

'0
'0
'0
'0

0'
FURTHER

STORAGE

PERIOD IN

MONTHS

FIG. ll.-Grnphs indicating percentage germination on moist filter-paper at 20 cC. and in

soil in t~ gr""nhouse, of onion seed.~ with moisture l"Ontents of 106 and 3"5 per cent held
for twenty-four months at 25C. and then pbced in paper seed-pad.,els or tin cans ror a further
storage: period of onc-half to si::l: months at Jocc. and 76 per cent relati,c h\1Ill.idity.

15

~hc.r~forc

bcen ahle to reuin more of their initi:ll gcrmiJl:ltion encrgy, "ere

;\bl~ to survive six months of packering, even when air-tight packctf; wer("
used. Low temperature will bring about similar eficcts when the moisl:\lfc
content of the seeds is unfavourable. Onion seeds with 10.6 per cent of
moisture which had been stored for f;ix or twenry-four months at Soc.
were much more rolerant of further storage in paper set..-d-packers than
those previously stored at 25"C. for the same length of time. This
Jifference was especially striking at the menty-four-monrh period.
Myers (I9{2) found that sealed lots of onion seeds stored in a refrigcr.ttor
will germinate well on removal to 'narural' conditions for three months,
while unsealed lots deteriorate rapidly.
A practiC31 application of sealing in tin cans has lx:en made by some
seed companies among'" bich is the Associated Seed Growers, Inc., of
New Haven, Connecticut, who have described their 'canned seeds' in a
monograph published in 195"" (Anon., 1954'/). They point out th:tr the
moisture content of the seed requires no !'>J}ecial adjustment if the seed
stock is in a region of low average relative humidity. They Ill:l.dc 'vigor
ralings' of stored seeds and concluded (Anon., I95-', p. 22) that <it is
feasible to preserve high initial viability and ";gor of vegetable seeds by
storage in moisture-proof sealed ems after the seed moisture content has
been adjusted to a safe lcvel which has been predetermined experimenr:ally.
Seed of onion, pepper, tomato, and certain other species is being Vigorpak
processed, i.e. conditioned and hermetically scaled, and offered under
the rCb,ristered name of "Asgrow Vigorpak seeds".' Some of the ap~rus
thcy use is shol\'fl in Plare 6, and germination of cucumher seeds stored
for tvm-and-a-halfye:trs in rin cans is pictured in Plate 7.
Air-tight tins werc also found the most satisfactory conuiners fur
cucumber, lettuce, C3trOt, cabbage, and bect sceds in Queensland,
Ausrr-.l1ia (F. B. Coleman & Peel, 1952).
From timc to time, so-called moisrure-proof plastic mat~rials have
appeared on me market. From these, water-resistant bags of various sizes
have been Ill:l.de to protect such m:nerials as foods and textiles, and parts
of machinery, from the hannful action of excess moistUre. Even though
some moisture does penetrate the plastics, they have served the purpose of
prolonging the usefulness of the encased materials in such places as me
humid tropics. Barton (I 949) tested several of these plastics to see whether
they would protect seeds from excessive moisrure. Eight different trpes
of polyethylene and \'inyl films allowed water to reach the seeds so that
ther could not be used for safe storage. Heat-scaled \-inyl laminated
aluminium foil, on me other hand, proved to be an effective moisture
barrier and could be used safely to store onion seeds under conditions of
high relative humidity. The importance of perfect scaling of the foil envelopes for safety in seed storage cannot be ovcr-cmphasizcd. If humid air
is allowed to enter through an imperfect seal, the seeds acquire \\ater
readily and deterioration is hastened.

106

--

EFFEcr OF PACKETS ON

S~]'J)

ViABILITY

When one sp<:aks of packets of seeds, the thought of small packets of

vegetable or flower seeds usually comes to mind. However, packcting
plays an important role in the life-span of other seeds, such as those of
coniferous trees, Seeds of Douglas fir (PselldotJllga taxi/oUa Britt.) and
Western hemlock (TJuga beteropbylla Sarg.) present a problem to the
seedsman and forestcr on at least three counts. First of all, frequent scedcrop failures make it difficult to maintain a constant supply of secus for
planting. Douglas fir trees, for example, produce a good crop every three
to seven yc..'lrs, with light crops in between. The setting of good seeds in
cones of Western hemlock is also likely to be sporadic. Another factor in
the availability of seeds of these twO species is their relatively short lifespan under ordinary conditions of storage (Heit & Eliason, 1940; Anon.,
1948). Still another problem, related to the second mentioned above, is
that of retention of viability under shipping conditions. All of these
difficulties arc magnified by the tremendous importance of dlcse trees,
especially Douglas fir, for timber, and the consequcnllarge demand for
their sccds.
It has been ShOlVTl in Chapter IV that sub~freezing storage is better
than temperanues above freezing for mainta.ining the viability of many
different kinds of seeds, including those of conifers. Furthermore, a
recent work (Barton, 1954) has shown that -18C. is better dIan - 1 J
or -4"C. for mainta.ining the viability of cerrain conifer seeds, including
the two species considered here.
Some seed companies now usc sub-freezing storage rooms, but the
question of the behaviour of the seeds upon removal [rom the cold storage
and packeting for shipment, and their ability to survive the shipment
itself, have been matters of concern. Also, some customers have been
reluctant to purchase so-called 'frozen' seeds.
We have already seen that the best condition for maintaining viability
of onion seeds, as measured by testing immediately upon removal from
storage, is the condition which permits the survival of the greatest number
of seeds during periods of subsequent storage. Detailed results of such
controllcd tests on Douglas fir and 'Vestern hemlock seeds are now
available (Barton, 195)
Seeds of Douglas fir and Western hemlock of the 1950 crop were
received in March 1951. They had been kept at oCoF. from the previous
December and were shipped to Yonkers, New York, by air express. Two
temperatures, SoC. and -J8C., were used for storage. Storage at S"c..
was in a large room with :J fairly dry atmosphere for the first two years
but with the atmosphere much more humid after that time. Sub-freezing,
-IB"C., was secured in a food freezer. Both temperatures wcrc constant,
varying only l "c. Seeds with two different moisture contents were used.
'Vhen received, the Douglas fir seeds contained 5.8 per cent of moisture,
calculated on the basis of the dry-weight of the seeds. Some samples were
stored without any moisture adjustment. In other samples, the moisture

'07

SEED I'RFSF.RVA1l0N ..\..1\ID LONGEVIIT

content was increased to 3,6 per cent by sprt.'ading the seeds in :l very
humid room at Soc., where they absorbed the desired amoum of water in
three days. They were then placed in a scaled container at - +"c. for one

week in order to ensure uniform distribution of the moisture increase

throughout all the seeds of the lot. Samples of the seeds with 5.8 per cem

of moisture were placed in canvas bags for storage at 5C. and -tSc.
Other samples of the same lot, as well as of the lot with the moisture
content increased to 13-6 pcr cent, ~ere stored at the same lcmper:ttures
after having been placed in tin cans with tight-fitting lids, further se:tled
with sealing wa.1:. Each sample weighed approximately 90 gm. and consisted
of about 8.400 seeds, one sample being intended for packeting testS after
each origi031 storage inter",al.
Moisture determinations made on seeds of Western hemlock upon
receipt indicated that they contained I I per cent. One lot was stored without any adjustment and another was dried to a moisture content of 7'7
per cent by spreading the seeds in the laboratory for four hours. Ori;irul
storage in c:mvas b:J.gs and sealed containers at Soc. and -18C was
as described above for Douglas fir.
All moisture determinations were made by drying the seeds in a
vacuum oven at 7SC. for forty-eight hours. Under sealed storage. the
initial moisture contents were maintained for the entire test period. The
seeds stored in canV':lS bags were subjected to moisture .fluc[u:l.lions
corresponding to the humidity of the surrounding atmosphere. At the
end of two years ofstorage, all lots in can..-as bags at both -18C. and Soc.
contained approxim:l.lely 13 per cent of moisture. Subsequentlr, the
atmosphere in the Soc. room became much more humid, resulting in a
moisture content of 22 to 26 per cent-in the seeds by the cnd of tlle third
year of storage. At -18C., the seeds still contained about 13 per cent of
moisture after three years.
Samples of both' species were removed from the original storage
conditions after 6, 12, 18, and.24 months and packeted in manila envelopes,
foil envelopes. and tin cans for further storage at 5 and 30C. Foil
cnvelopes were made of vinyl-coated aluminium foil sealed by heat. Tin
cans were sealed by tight-fitting lids and sealing wax.
Storage of the 5C. packets was in a large room. the atmosphere of
which was dry at the be<;inning of the test. '''hen the atmosphere of tltis
room became humid at the end of two years, all the Soc. packets were
moved to a small, dry chamber maintained at this temperature. Storage of
packets at 30C. was in a dr)' room with good air circulation, resulting in
a low moisture cOlHcnt(about ";'5 percent)ofthesccdsin manila envelopcs.
Tests of the germination capacity of the seeds were made at the time
of removal from the original storage conditions for packeting, and after
0'5, r, 2, 3. 6,12 and, in some cases, 18 and 2~ months in the packets.
The results of these tests have demonstrated once again that subfreezing temperatures are to be preferred to aboyc-freezing ones for the

'08

--

EFFECT OF PACKETS ON SllD VLillILITY

maintenance of high quality conifer seeds. It has been shown furtllcr that
the better the original storage condition, the greater will be the value of
the seeds when thcy arc rcmO\'ed from storage for planting or packeting or
shipmcnt and, ultimately, germination. This invalidates objections to thc
use of 'frozen' seed. Experimental results have shown that Douglas fir
and \Vestern hemlock seeds stored at -ISC. for as long as two years
retain their full germination capacity and do not suffer, by virtue of their
storage, upon transfer to a higher temperature. On the contrary, they arc
more resistant to subsequent deterioration than seeds which have been
stored above the freezing-point, and which will have lost some of their
germination vigour. For example, Douglas fir seeds packeted in manila
envelopes held at 30"C. after SL,< months of origi.n.1l opcn storage, deteriorated at the same rate whether the original storage temperature was - ISO
or Soc. The same relationship held after twelve months of original
storage, though the actual germination percentages were somewhat
reduced. After twenty-four months of original storage, however, the
great decrease in germination which took place during storage at Soc.
made the seeds worthless for packeting, whereas those stored originally at
-lSC. were still of high quality. In other words, any condition which
prevents deterioration of the seeds in storage, whether that condition be
sub-freezing temperature, reduced moisture content, or both, permits an
increased delay in planting. Some seeds held at a temperature as low as
Soc., but in high humidity, are worthless after a few months, whereas
seeds of the same lots stored at sub-freezing temperatures for years on
end may be as good as fresh seeds in every way.
Six months oforiginal storage at - I8 c C. plus t\\'elve months in packets,
or twelve months of original storage at - 18C. plus six months in packets,
had the same effect on subsequent germination of Douglas fir seeds when
the packets were stored at Soc. Furthermore, the conditions were as
favourable for retention ofviability as an original storage period of eighteen
months at -18C. 'Vhen the packets were stored at 30C., however,
there was a greater retention ofgermination capacity after twelve months of
original storage at _18C. plus six months in packets, than there was after
c
SL-":: months of original storage at -I8 C. plus t\,-e1ve months in packets.
The survival of seeds upon removal from original storage and packeting,
then, is directly related to the time they arc exposed to an unfavourable
temperature or humidity-whether that time be during the original
storage period or after packeting for sale or shipment.

<09

XlI
METHODS OF TESTING FOR VIABILITY
AcruAL germination of seeds in soil in the greenhouse or field, or in some

suitable medium in the laboratory in controlled-temperature chambers,

has been tbe tried :md oue method for determining germination C3.p:1city_
However. this is not always as simple as it might appear. In tbe first place,

there is apt to be a discrepancy between the ability of seeds to produce a

root under controlled conditions on a medium such as filter-paper or a
moist paper towel, or c\-en in sand, and the actual srand of seedlings
produced in the field (Goss, 1931; Kjaer. 19+5; and others). Some of these

relations have been worked out, so that the field performance can be
forecast from the results of a laboratory lest. In the second place, many
seeds have special requirements for germination, and obviously \\'ill Dot
produce seedlings unless these requirements are met. For non-dormanr
seeds, this may involve a special tcmperature, exposurc to light, or
mechanical or chemical treatment of the seed-coo.ts to make them permeable. For dormant seeds, it may mean an after-ripening period either
in dry storage, such as is characteristic of certain grains as well as of
lettuce and some other seeds when freshly han'estcd, or in a moist medium
at a low temperature, which is a requirement of the seeds of many trees
and shrubs.
Not only may there be special requirements for germination, but thesc
may ehangc with storage. For example, low-temperarure pre-treatment is
reponcd to be more effective for old Pinus pa/uslris seeds than for fresh
ones (Barton, 1930), and the germination of old ~rden seeds is claimed
to be benefited by soaking in ethylene chlorhydrin (Ruge, 1952). Many
seeds having specific germination requirements at harvest time will
germinate over a wide range of conditions after dry storage for a few
months. For general discussions of these special requirements for germination in seeds, the reader is referred to five general articles (Barton,
1931}d, 1953; Barton & Crocker, 1948; Crocker, 1948; Crod:er & Barton,
1953). Some ofmese special requirements have been described in sections
of the present book dealing with specific seeds.
Even when no special treatment is necessary to bring about germination, it is essential tbat there be a smndard method for testing seeds of
commercial importance, in order that they may be tested or shipped to
any place in the world. This has led to the de.,etopment of inlernational
rules for seed testing. These rules arc formulated and approYed by the

"0

l\-1Ell-l0DS OF TISTING FOR VIABILITY

=
-

Internation;ll Seed Testing Association and are revised as found necessary

from time to time. The latcst of thcse is to be found ill the Proceedings of
the International Seed Testing Association for 1956 (Anon., 1956). These
rules were adopted by a meeting of the Association in Dublin on 29 May
1953, and were revised at the Paris meeting on 8 June 1956. The original
rules came into force on 1 January 1954,for the Southern Hemisphere,and
on 1 July 1954, for the Northern Hemisphere. The amendments became
effective upon adoption. Hence the earlier International Rules for Seed
Testing were cancelled.
Not only arc there international rules, but individual countries also
have seed acts governing the testing and sale of seeds within their
territories. These methods as applied to the United States have been set
forth in a Manual for Testing Agricultural and Vegetable Seeds (Anon.,
1952a). J'vluch research by manr inve!>1:igators has preceded the establishment of such rules, but this work need not be described in the present
volume.
Special requirements for the germination of certain seeds have not
been met by some workers testing viability after storage under various
conditions. For this reason, the seeds have been reponed non-viable when,
in fact, they ,yere capable of germination. This has applied especially to
seeds with impermeable seed-coats or dormant embryos. Germination of
elm seeds, for e.xample, is inhibited by temperatures above 25C. when
they arc mixed with moist granulated peat-moss, and very few seedlings
are produced in soil from fully viable seeds wruch have not been pretreated. Pre-soaking in wmer for twenty-four hours, as well as pretreatment in moist granulated peat-moss at 5"C. for one month, resulted
in greatly increased seedling stands in soil in the greenhouse (Barton,
1939tl). Unless these facts arc l-no\\"l1, it is apt to he assumed that the
seeds, which are presuma.bly short-lived, are non-viable. The life-span of
elm seeds was discussed in Chapter IX.
Germination tests of conifer seeds are aLc;o often inadequate and do not
express their true viability (Mirov, 1936). Furthermore, it may happen
that types ofseed which germinate easily without pre-treaonent when they
are fresh or when relaining full viability in storage, may require prctrcatment for germination of deteriorating samples. This has been
demonstrated for several secds, among which are those of the longleaf
pine, Pinus palus/Tis JVlill. (Barton, 1935a). From a. given storage condition,
these seeds retained only half of their original germination capacity as
measured from plantings made directly in soil in the greenhouse, but if
the same seeds were pre-treated for one or two months in a moist medium
at SoC. before planting in the greenhouse, four-fifths of the original
germination percentage was obtained. Also, treatment with chemicals to
prevent attacks by micro-orb,ranisms has been reponed in many' instances
to increase the seedling stand from old seeds to a greater extent than from
fresh secus (see, for cxample, Wallen e1 til., 1955).
m

SEED PRESERVATION

.oU~l)

LONGEVITY

QUICK VIABILIW TESTS

Bcca.usc of the special treatments necessary, anJ the collSequCllt

delay, in finding the germination capacity of seed-lots of vital importance
to the seed tradc and to planters, an interest developed in \\'.1 ys and means
of estimating the performance of the seeds without the ne<:essity of
germination tests. Perhaps the earliest attempt to get a rapid estimation
of the germination capacity of seeds was through the 'cutting' test. The
seeds were simply cut open and the number of empty ones recorded. This
gives a fairly accurate measure of germination expectancy when the seeds
are fresh or of high quality (Barton, 1930; Viado, 1938), but permits no
evaluation of deterioration in the embryo.
A variation in the cutting method ,vas reported by Dorph-Petersen
(1925) and used by Trautwein & Wassermann (1929) and Weisenfeld
(1929). Used for cereals and known as the 'half-grain' method, it involved
cutting off one end of the seed so as not to injure the embryo. -The remaining portions of the seeds were then placed on a moist substrate where the
cotyledons turned green within a few days if the embryos were viable.
Zachariew (1939) used this method as late as 1939 for seeds of Pillus
sylvesfris L. An extra refinement was made when the seeds were fully
excised before testing.
EXCISED E,"IBRYO TEQU'iIQUE

Because of the continued dormancy ofthe embryo in dry storage, many

tree and shrub seeds require a period of from one to s~veral months in a
moist medium at low temperature. where they after-ripen, before germination performance at higher _temperarures can be ascertained. Such
delays in the determination of viability may cause considerable inconvenience as well as increase the consumer's risk of bUjing seed stocks
whieb.have not been tested. A rapid means of detennining the germination
capacity of such seeds has, howeYer, been developed in the excised embryo
technique.
As early as 1906, Crocker emphasized the importance of the seed-coat
in germinating weed seeds, and pointed out that removal of the eoa.ts of
some weed seeds removed the germination barrier so that results could
be obtained within a short time. Harrington & Hite (1923) removed the
coats from dormant apple seeds and found that, if they were kept moist,
the cotyledons became green and some of the radic1es started to grow.
Flemion (1936) and Tukey & Barrett (1936) were the first to apply this
method to a rapid determination of the germination power of dormant
seeds, and hence to use it as a measure of the viability of such seeds.
Working independently, these authors reported on the use of the method
for determining the germination capacity of pcach seed. The rapid
technique as llsed by Flemion for many IYpes of dormant seeds (Flemion,
1936, 1938, 1941, 1948) compared favourably with germination tests
which followed after-ripening: at a low temperature.
IT2

'\'iETHQDS OF TESTING FOR VTABIUTI'

The general method which Flemion used consisted of removing the

hard outer CO:.lt, aftt:r which the seeds were soaked overnight in the laburatory, before removing the thin inner coats. The 'excised' embryos "wcre
then placed on a moist surface, usually filter-paper, in diffuse light in the
laboratory, where the greening of the cotyledons and extension of the
radicle indicated viability. It is not necessary to place excised embryos on
a nutrient substrate, as recommended by Tukey & Barrett (1936), in
order to test their viability. If the embryos have lost their viability they
fail to turn green, and will rot within a few days. Plate 8, taken from
Flemion (1938), pictures the appearance of excised embryos of different
germination capacities. Flcmion (1948) has summarized the work done
with the excised embryo method and has listed the species that have been
tested, together "';th literature references. This summary is given in
Table XIV. The original literature citation numbers have been changed
to authors and datcs in the table hcre presented. In addition to those listed,
Treccarn (1951) used the Flemion method for testing fruit-tree seeds.
A comparison of [he excised embryo and the terrazolium methods of
testing for viability will be given in Chapter XIII.

TAlJLE XIV
LIST OF SPECIES, WITH LITER.'l.TURE REFERThO:S, '\VHIO-l HAVE
TESTED BY THE RAPID VIABILITY METHOD

I'lant family

BEEN

Reference under litcrarnre

cited

Species

Amar.ulLhaL"ele
BerbcricbLtlc
Bignoniaceac

Heit (19.B)
COlllpllf/m{l g/obOIa
Heir & Nelson (19+t), Heir (t943)
Bahtris thunbergii
BigllolJia (Campsis) radicalls Flcmion (t9.P), Heir & Nelson

BOt:lginaceae

C)'/Iog/(JssuIJI i/maPilt
Symphoriwrpos racemosus
Celas/rus scani/ens
EU(JIlYIIlJiS sp.
C(Ju(Jpsis
Wyethia scahra
Comus sp.
Cucurbita, Cucumis, and
Citrul/us !>pp.
Echillllcystis
Shepherdia argenlea
Ellcommia /llmoidu
Euplwrbia margillata
Liql/l-dambar styracifiUQ

Caprifoliaceae
Cdtstrn=e
Composit:lc
Comaceae
Cucurbitace-Jc
Elaca!!I1:Jceac
o
.
EUCQrnmlaceae
Euphorbiaceae
Ham:unelidaccac

Hamamelis spp.
Lq;uminosac
Lythnl.ceae
l'vIa!,'1lo1iaCdC
I\ohrtyniaceac
H

Caci! canaatllsjs
Lagerstrwl/lja illdica
Aragl/olia ruumillilta
MarlY'lia

"3

(t9-l-1), Heir (1943)

Heir (194-3)
Flernion (t941)
Heir & Nelson (19+1), Heit (19-1-3)
Heir ('9.B)
Heir (19..1-7)
Flcmion (1941)
Heir (r943)
Heit (1943)
Heir (19.1-7a)
F1crnian (19-1-1)
Flernian (1941)
Heir (H)-I-3), Heir ':fj3a)
Anon. (194U),
cit & Nelson
(1941), Heit (1943)
Flcmion (1938), Heit & NeL~on
(1941), Heir (t9-1-3)
Flernian (19-1-1)
. Flemian (1941)
Heir (U}j.3)
Hcit& Nelson (t94T), Heil (H)-H)

TABLE XIV-Conrd.
Pi:lnt family

Oleaceae

Spedes

Fraxinlls Spfl.
Chionon/fms spp.

Pinaceae

Mmodora-spp.
Abies cOllcolor
Pillus spp. (13 species)

ROS:Iccac

Psrodolsuga-Sp. (Douglas
fi,)
Tsugo caNadellsis
Cluzroomdes (Cyao/lia)
japOllica
Cralaegus spp.

LYOI/otflalllnu.l jlorilll/lllills
lHal1lS spp. Apple
PrIll/US spp. (9 species)
Apricot
CherTy

PC2ch

P'"",

Rhodo/ypos J:l'rril}it!t'$
Rosa spp.
Sorbus alauporia
Spargalli.aCC;1C
Urnbcllicre
Urticaccae
Vcrbenaceae

Spar/:aniulII euryc(upUIIJ
TrQchYfflcnc (Didiulls)
coC/ulco
COllnobis sativa
Ulmus omuuol/o
C((lIicarpo purpllrm

Rc[..,rellf:e under iiter-Hure

cited

Anon. (r94Ic), Heit& Nelson (19-\ I).

I-kil (1943). G. 1'. Steinbauer
(19-\4)
Flernioll (1938), Anon. (194"),
Flcmion (Iy.p), Heir (194])
Flcnuon (19..j.l)
Hcit (19..j.])
Flemion (1938), Baldwin & .Flemion
(J9P), Flemion (19.P), Heit
(194J)
Flemion (1938), Flemion (19.P)
Heir (1943)
AnOll_ (19-\1C). FlClIl.ion (1941). Heir
& Nelson (19..j.1), Heir (1943)
Flemion (1938), Anon. (19'pc),
Flcmion (19..j.I), Heir & Net~on
(1941), Heir (19..j.3)
Flcmjon (19.P)
Flcmioll (1938), Heir (T9.0), Heil
(t946)
Heir (1946)
Flemion (19P), Heir (1943), Heir
(1946)
Flcmion ('936), Heir & l\'elson
(I94 I), Heit (19_B). Tuke)' (1944),
Heit & Legnini (1945), Heit(/9,!-6)
Flcmion (1941), Heir & Nelson
(T9-1 1 ), Heir (1946)
Flcmion (HUS), nCnUon (T94 J ),
Heir (1946)
}'lcmion (1938)
Heit (T943)
Flemion (1938), Anon. (H)-fU),
Flcrnion (1941), Heir (1943)
G. P. Steinbauer (19-14)

Hci[ (1943), Heit (I947a )

"''1emion (1938)
Barton (193ga)
FJcmion (J9.P)

A variation of this technique was used by Krstitch (195 1-5 ) for

2
embryos of PiIlUS nigra. They were pbeed on gelatin containing tannin, in
which circumstances good embryos grow and turn reu in twcmT-four to
forty-eight hoUT!> at 23 to 25C. but dead or weak embryos haw a
brownish colour and arc not ttl1'gid.
~I'vfore recently Heir (1955) has described the embryo excision method
as 'efficient, accurate and timely' for seed analysts. The seeds he tcsted
included fifteen species of flowers and vL"getables, twenty species ofshrubs
and vines, seven species of fruit trees, and forty-seven species ofconiferous

"4

Ph~f~.

hy collrusy ASf(/{;llld Sud GrOIr'I!r!, Inc., ."'ap Ha'C'nI, COllnnlhul, US.A.

Pun: 7
Germinating cucumber 5ttd-lots of diffcrent rnDisrure levels from lin cans hermetically
!;Calcd and ~torcd at three conStant remperarnrcs for tv.o-and-a_half years. ricmres taken at
rime of first counts. From some 10m, a few dead mouldy seeds ha,~ been rernon~d. Germinations indiaHed in the table gi'cn below are [()tal~.,r first 3nd fin3! counts, and arc al'engcs
of four replic:ucs of 100 ~eed~ each. They nmge in columns from left to right in the illutr:Ition. Vigour r:ltings: E, excellent; G. good; F, fair; D. scrd dead. Germin3tion befor~
storage "105 98 per cent, and the sc<:dting vigour was E. The results after two-and-a-half
years follow:
Storage
Germ.
"luist, Germ.
.Moist. G=. Vigour
%
Vigour
temp.
% Vigour %
%
%
60F.
,8
E
8,
G
E
7'9
96
't...
E
go
86
70 1'".
G
o
D
93
90F.
F
8'3
0
D
E
5'3
95
(From Anon., (9548)

Ii

PLATE 8

Development of excised embryo~ On mo~~t filter-paper in Petri dishes at room tempcraturc:

K, natural size; E, M, and;.; x -1-; all others x 2. (Rcading from left to right in each cas!:.)
A and B. RhQdQtypQ, k(m-I)idu. .\, Freshly nrued emhT)-o and ivc-day dnelopmcnt on H)
November, 193/, ofllie 1932, 193-1-, 1935, 193u,md 193/ (2 tmbr!os).::rops, respcdi,d!. Not~ bd:
of growth in lypiCll embryo from 193::lcroP, limited dcvelopment in '933 crop, and increasing ngour
in '935, 1936 and 1937 crops. Il. Embf)'os of 1932 amI 193-1-croPS after Ien days. The '932 embryo
is Wldergoing de~lru<:lion by organisms and the 19H emhryu is stilll'Ul1Sider~bly behind the lh-ed1y deYclopmcnt of embryos from bter lTO~ as shown in A. The 19_H embryos han' a pale ~-dlow
grecn colour, rardr cxhibit any hrpocotyl del"l:lopmcnt, and dcteriorate rapidly. Embryos which
~how this fype of dCYclopmcnt arc low in I-igour, and only a low perccn!age of seedlings is obtained
from them when the intact seeds are after_ripened :H a low temperature.
C, Applc crnb~-os. First, a freshJy-excised embryo is shown and thcn (after six days on moisf
filter-paper) there arc Iwo non-,--iable ones followed b~' fwo ,--iahle cmbr~-os ~howing (ypiall greening
on lTItyJedons and new gro\nh. Photographed 9 ~o\-ember, 193/
v, Cra/{ugus mJ$~gal!i. Ten-dar del-dopmen! on I Febma~', '938. The m-st three ill\lstrations
arc of emblJ'os of the 1928, 1930, and 1933 crops, reslX'Cti,-cly. The embryo of the 19z11 crop has
undergone marked deterioration while the emhryos of the 1930 and 1933 crops arc also shown to be
nun~,'iabk .\t the ~"me time the behaYiour of two \iable embT)'os is shown...h the eXLreOle right
illll is an emhr~'oora 'eed ofm" 1936lTOP whi<:h had been in moist peat_mns' at 5~C. for one year.
Obyiously the intact seed lud not afrcr-ripcncd during this time bllt had remained .iable_ The
viable embfJ'O of the 1935 crop (second from right) \lill, \lith time. also show similar enlarl,(cmcnt
of the cot~'kdon next to the moist filter-paper. It is n... dil~- seen that "iable hawthorn embryos do
not exhibit thc extremely rapid dnelopment obtllined with ,-iable Rhodol)'pos seeds_
, Sorbm auwparia cmbryos. On the left is a photot,'T:lph of a frcshly--c"ci~cd embryo, after
which follows a dead embr~'o of the 1930 crop and a viable embryo of the 1937 crop after SIX lbys
(18 Noyember, 193/) on moist filter-paper.
F. Here arc shown (18 "'ol'ember, 1937) embfJ'05 of Witch-hazel of the 1934, 1935, and 1936
crops, respectively, after th-c thrys. The 1934 embryos are dead and are undergoing deterioration,
the 1935 embfJ-oS which arc low in \-iu1itycxhibit Yc.r~little acti'ity-thc. cotyledons merely spread
apan-while the \iable Hj36 embryos arc growing. In G, H, and J arc shown in each lllSe two
typical embT)'OS of the '9H and 1936 crops, the older embryo appearing on the left. The cmh~-o,
in G have been soaked for twemy hours in a s.arurated solution of para-ilinitrohcnzcne followed by
one-half hour in dilute ammonia, and those in H hayc been treated for twenty hours in I per cent
pota,~ium tellurite. J illustrated the beha,iour of embryos after live day, on moist fIlter-paper.
The chemical tests gh-e some oolollr e,-en in the 193-1- crop, :md reactions intermediatc bern-cen
those shown al,o occur, so that it is difficult in man)- Cl.'lCS to doxide by chemillll tests whi<:h
embr~'os arc al"e. The results Oil moist fillet-pap<., arc much easier to intcrprd.
K. Prllnus amairono embryos :u"tcr se,-cnl \I-eeks on moist filter-paper. All these embryos arc
viable and the ""rious types of dndopmeol are illustrated. The dead (nol shown here) ernbfJ'OS
deteriorate rapidly' while the ,-iable ones develop slowly.
L. Fringe-tree (ChiollalIrhus "irgillk,.). The embryo on the left illustrates the deterioration of a
non-viable embryo after fOllf days, and the other photographs illustrate a .-iable embrJo after four,
six, and tWCOT)' days, respectiYelr.
11.-1. Douglas Fir. A freshly---excised embryo appears On the left, then vi"ble and non_viable
embf)-os after four days on moist filter-paper.
N. Pinus rigida. A freshlr--exeiscd embf)-o is seen on the left, and then a non-\-iab!e embryo and
twe viable embtyos crlUbitmg 10"- amI high ,irality, respectively, after eight da~-s.

PUTt 8

METHODS OF Tf.5TING FOR VIABTJ .TTY

--

:lnJ bro:vl-Ieafed trees. Nord (1956) used 400 excised embryos, in lots of
lOO each, for viabilit)' test of bitterbrush (Purs!J;f/ fridel/lfi/If) seeds.
Altogether, it appears that the excised embryo method is reliable and
capable of morc exact interpretation than staining tests. However, it
apparently has not been included in any official, international seed testing

rules.
PLASMOLYSiS

Plasmolysis, or the shrinking of the protoplasm from the cdl~wal1 in

the presence of solutions such as a saturated sugar solution or a 2-00rm:11
potassium nitrate solutioIl, has been recommended for the determination
of germination capacity (Doroshcnko, 1937; Niethammer, 1942). But
it has not been extensively used, owing perhaps to the skill required in
preparation and in the microscopic determination of the germination
values.
ENZYME ACnVITY

With the realization of the need for rapid viability tests of seeds there
came the idea of measuring their metabolic activity through some
particular process such as respiration (Qyam, T<)06) or the amount of heat
given off when the seeds were placed under germination conditions
(Darsie et (11., 1914). Biochemical assays also came in for their share of
attention. One of the first, and the most widely tested, of these was
enzyme activity. Much of the work in this field has been done on the
relatiOn of catalase activity to viability in seeds.
Crocker & Harrington (1918) were not able to use catalase activity as a
measure of viability of seeds of Amaral/Jlms and Johnson grass (Holcus
lJ(jfepemi.~ L.). Nemec & Duchon (1922, 1923), however, found a close
relation between germination c.apacity and catalase activity of Oats from
the crops of 189T to 1912. Peas responded in a similar fashion. Niethammer
(193W) and Brodskis (1949) found a general high correlation between
high catalase activity and high germination values. Lcggatt (1929-30), in
an extensive survey, found the catalase assay method promising. A
statistical analysis of his data, made later (Leggatt, 1933), showed that
viability, in the case of wheat grain, can be estimated fairly closely from a
determination of total and thermostable catalase, but caution was advised
on the routine use of such a method. These early reports are typical of the
differences of opin.ion as to the value of the catalase-determmatioll method
of measuring germination capacity.
W. E. Davis (I925) pointed out that dry seeds, such as were used by
previous workers, were not suitable for catalase measurements, as the
enzyme does not necessarily disintegrate with the loss of viability but may
remain unchanged for many years after the seeds become incapable of
germinatin.g. He postulated that catalase differences in non-viable and
fresh seeds would be accentuated by placing the seeds under conditions
115

SF.ED PRfSERVAT10N ..\.l"1) LONGEVITY

which ,,-ould initiate germination. lIe, therefore, soaked seeds in warm

water at a temperature high enough to inactivate c:l.tabse, but not high
enough to affect seed viability. Lettuce seeds s()aked in water overnight
at 32 "c. showed reduced catalase activity in dead or partially deteriorated
seeds, with no change in the amvunt of the enzyme in viablt:
. . caralasc activity of soaked seeds
I
- fd
d ' rather than
seeds. The catalase ranD, I.e.
cata ase activity 0 ry see s
the actual caralase content, thus becomes the imporram consideration in detennini.llg viability. He noted, however, a tendency for the
catalase of viable seeds w increase because of increased respiration
following absorption of water. The amount of oxygen in the water llsed
for soaking is also a factor in determining the amount of catalase activity.
The conclusion from this work was that catalase activity could he an
indication of viable as opposed to dead seeds in a sample, but could not be
used to estimate different degrees of deterioration. It could be of value in
the estimation of viability of dormant embryos in cases where disparch is
necessary. It could also be used to confirm re.'illirs from other rests or to
derect improper germination methods.
Takiguchi (1932) arri\'ed at similar conclusions from a comparison of
the catalase content of dried seeds and seeds soaked according ro the
Davis method. The 'catalase ratio' method of Davis was a1<;0 used by
Spacth (1936) for tree seeds and by B. N. Singh et ai. (1938) for seeds of
Zea- mays, Pimm (/fVense, P. sativum, Cajanus indicus, Triticum ;;ulgar",
and Gieer ([riettlm, with the general result that ratios close to or aboye 10
indicare viability and ratios lower than 10 will indicate loss of germination
capacity. However, as predicted by "\V. E. Davis (r925), decre.1se in
catalase ratio was not always proportional ro decrease in viability of thn:c
seeds at intermediate stages of seedling vigour.
The absolute amount of catalase in the resting seed of pine, in relation
ro the increasing amounts produced in seedlings within ten days of the
commencement of germination, should be used to indicate the final
percentage capable of germination, according to '\T. Schmidt (1929).
Nuccorini (1930) agreed with this modification of the Davis method.
Baldwin (1935) found that a few days' stimulation of conifer sccds in a
Jacobsen germtrultor gave sufficient differences in catalase activity from
the dry seeds to permit estimation of germination capacity. In this case,
the ratio behveell the oxygen evolved by the germinating seeds and the
oxygen evohed by the dry resting seeds, correlated well with actual
germination tests. Q!1otienrs greater than unity indicated seeds of high
qllality.
Ir has been reporled thar catalase measurements are useful in the
derermination of damage to seeds by X-ray treatments (E. N. Singh, 19..p)
and by soaking in solutions of phenyl propionic acid (Nanda, 1950). Al<;o,
successful use of the method has been cla.imed by Pcrersohn (1926) to
derect heat-killed white cabbagc seeds as an adulterant in expensive red

,,6

METHODS OF TESTING FOR VIABILITY

ctbbage seeds. However, most of the people who have used tllls test have
some reservations about its value in general use. Failure to get any
(.urre1:J.tion between germination and catalase measurements has usually
resulted from the use of dry seeds alone (Gracanin, 1927; Grisch &
Koblet, 1931; Knecht, 1931; Nazarova, 1937).
Other enzymes have also been shmm to indicate viability, but mostly
in connection with the staining of the living tissue. TIlls will be discussed
beluw. Direct measurement of oxidase content of cotton-seeds revealed
that it controlled both the speed of germination and the water absorption
capacity of these seeds (Nakatonll, 1936). McHargue (1920) reported a
correlation between peroxidase activity and the viability of seeds of
twenty species.
U-IE.MlCAL TRE..-\1J\IB\'TS

External application of chemicals which bring about observable

changes within the Jiving tissues of the sl-'Cds was used at an early date.
Perhaps the first was sulphuric acid applied on the cut surfaces of grain
seeds, resulting in the formation of a deep rose-colour in living seeds
v..ithin five minutes, while fifteen minutes were required for the appear~
ance of the colour in seeds of poor quality (Dimitricwicz, 1876). Lesage
(r922) used potassium hydroxide solutions for determining the germination capacity of seeds of Lepidium sarivu11I within four hours.
DYES

Neljubow (1925) mentions previous methods of testing germination

capacity without germination tests, including the measurement of carbon
dioxide production by secds which have imbibcd \vater (Qyam, 1906), the
method ofLcsage (1922), and the catalase method; but he states that none
of these was effective. Nc1jubow proposed that organic dyes be used. He
found that indigo carmine and acid violet, as well as a mi......ture of neutral
red and methylene blue, would stain dead embryos or dead parts of
embryos, but would not stain living embryos or living parts of embryos.
Viability as determined by these dyes agreed very well with the results of
germination te.<;ts. Issatchenko (1931) confirmed these results, emphasizing
that the embryos must be taken out of the fruit- or seed-coats before they
are tested.
The indigo-carmine method has been found effective for seeds of
Umbclliferae (Doroshenko, 1933), Caragalla arboresuns, Pinus sylvestris,
and Picea excelsa (Shefer-Safono\'"1l cf al., 1934), Aleurifes (TskoiabC, 1936),
and Pinus syivestris, Picea abies, Pseudotsuga taxifllia, and Abies alba
(Hao, 1939). Grimm ct ai. (1928) described the method as very simple a.nd
used it in numerous trials. The seeds were soa.ked in water for one to three
hOUTS, until the outer envelope could be easily detached with forceps.
After remonl of the coats, the seeds were soaked in the I : 2,000 indigocarmine solution for four hours a.t 26 to z8C. They were then washed

"7

SEED PRESERV.o\TION .-\.I\ID LONGEVITY

and their germinative power determined from the colour: seeds that
becamc entirel)' coloured were incapable of germination, while those that
remained uncoloured were capable of germination and yielded normal
seedlings. Partl)' coloured seeds, including those coloured at one end or
showing spots at both ends, had defective embryos and their sun-ival :l.fter
gennination could be predicted by the extent of the staining.
Bismark bro'i\TI is the best indicator for barley seed viability according
to Kornfeld (1930) who found that, with it, all non-viable seeds became
dark brownish-green while viable seeds showed scared)' a trace of the
colour. Germination e:~. pcriments with the grains which had become
coloured gave no seedlings. Methylene blue used previously to indigo
carmine (Turcsson,1922),and malachite green (Gadd, 1944), are other d)'es
which have been used successfully. A more complicated method of
suining was used br Tanashev (1938), who combined cresol red with other
chemicals such as phenolphthalein and xylenol blue. He nOted clut this
improved the dre-indicator method. Niethammer (1931), using intact
seeds of grasses and leguminous plants, obtained good tests for viabilit),
with methylene blue, neutral red, and orange G. The permeability of the
seed-coats to the dre is a nul factor in the usc of this method. Also, the
permeability of some coats for specific dres may change as the seed ages.
Perhaps it was for this reason that tests 'with 01 per cent Congo red did
not always show a good parallel between staining and germinability.
SElL-':IU~1 .~'\'D

TE:U.URIUM SALTS

Sak:1ta (1933) objected to the indigo--carmine method of Ncljubow

on the grounds that if the seeds were left in the solution too long, viable as
weU as dead seeds became coloured. He used teUurium salts instead.
Although some other workers ha.e used tellurium salts, the majority of
them have used salts of selenium, which give essentially similar results.
The method has been described by Eidmann (1938). Seeds from which
the outer coating has been rema.ed are soaked in water for twenty-four
hours at room temperature and then further soaked in a 2 per cent
solution of soJium biselenite (NaHSc0 3 ), adjusted to pH 4.5-5.0, for
fortr-eight hours. in contrast to the dyes which stain the dead tissue, the
selenium compound stains the living tissue a permanent red by depositing
elemental selenium. The embryonic tissue of dormant as well as nondormant seeds is affected. Dead tissue remains uncoloured. The intensity
of the coloration increases in proportion to the degree of vitality of the
seed. This poses the question of judgemenr- of the results and requires
experience to be able to interpret the findings successfully. In general,
selenium tests of good seeds give reliable results, but seeds of poor quality
give variable results which do not agree closely '\~th germination tests in
the case of some conifer seeus (Hao, 19390).
L. P. V. Johnson (1947), as a result of his study ofthe use of the sodium
biselcnite method on seeds of a wide range of agricultural plants,

uS

1
:

ME'!l IODS OF TESTiNG l'OR VLWILITY

concluded that it was generally satisfactory provided the pre-soaking step

W.IS modified to include sixtccn hours of pre-soaking and eight hours of
aeration, He pointed out, however, that the relation between the biselcnite reaction on one hand and seed germimbility and seedling vitality on
the other, must be established for each species before the results can be
interpreted correctly. Thc method, suitably adjusted, could be used for
evaluation of the seedling performance which could be expected from a
given seed-lot. In the case of seeds which have been killed by Fusarium
infection or by frost, the selenite mcthod gives results \vhich are too high,
according to Gadd & Kjaer (1940), Lakon, who might be called the
champion of the tetrazolium method to be described in the next chapter,
also found rhat the selenium method was quicker than the germination
method, gave the highest germination capacity, and was not affected by
dormancy (Lakon, I940).
The selenite method has also been found useful for tests of seeds of
Citrus (Monselise, I953), winter barley (Eggebrecht & llethmann, 1939),
and Karoo bush (I-Ienrici, 1943).
Boiled, ground, mashed and finely cut seeds or embryos of Pinus
nigra, Picca abies, and Fagus sylvatica, and boiled and unboiled expressed
juices of these seeds, were not coloured by sodium biselenite, but the
undamaged living seeds were coloured red (Wach, 1942), The living tissue
is evidently required for the reaction. Plaut & Gabrieiit-Gelmond (1949)
presented a table of results of what they call the two most promising of
the staining methods for te-<>ring viability, namely selenium or tellurium
salts and 2,3,5-trtphenylterrawlium chloride. The table lists a considerable'
number of cerC:ll, forage,and vegetable-crop as well as conifer seeds, which
have been found to respond to one or both of these treannents. These
authors believe that the results warrant further intem,ive smdy for
routine testing procedures. Telluritcs ,,,ere considered more reliable than
staining with methylene blue, catalase activity, or the presence of soluble
sugars, for detcrmining the vitality of seeds (Toniola, 1949).
DIN1TROBE:.~ZOL

=
=

Gurewitseh (1935) developed a method of detcrmining viability,

which was based on the power of the living cell to reduce dinitrobenzol
through respiration. The reduction products, nitrophenylhydroxylamine
and nitranilin, are distributed differently in the tissues of the seed, as is
indicated by the unequal intensity in the oxidation-reduction process.
When ammonia is added to viable seeds, nirrophenylhydroxylamine gives
a characteristic purple colour. The uriginal tests were used successfully
with wheat and rye grain, stained by solutiuns of meta-dinitrobellZol.
Later, however, Ebes (1936) and Gurewitsch (1937) determined that the
effects were due to the orflzo and para isomers present as impurities in the
me/a compound, and that the latter alone was ineffective. \Veise (1937)
confirmed the work of G urewitsch with wheat and rye grains and extended

"9

SEED PRESERVATION Al\'D LONGEVITY

the method to beans and peas. It could not be used with asparagus or Iruittree seeds.
RESAZURTN

The use of resazurin for viabilit)T tests has been developed at Rehovot,
Israel, by Plaut and his co-workers (Plaut & Halfon, 1954; Plaut & HellerCohen, 1956; Plaut el a!., 1957)' Resazurin is blue in alkaline solution and
red in an acid solution. It can be readilr reduced to a red colour, followed
by orange and finally white. A solution of 0.003 per cent rCSazurin was
used for testing the seed of pea, bean, and cucumber (plaut & Halfon,
1954) The embryos of viable seeds remained white, but those of nonviable seeds stained red or blue. In this concentration, the chemical docs
not kill the embryo, so that the same seeds could be tested for germination. It was found that all the seeds with white embryos germinated, while
those in which the embryos stained blue did not. Tests on cereals were
performed by removing the periearp and immersing the grains in a O'O!
per cent resazurin solutiun for SL'{ hours (Plaut & Heller-Cohen, 195 6).
A blue stain of the coleoptile or of a major part of the germ (embryo)
indieated that the seed was dead, whereas a red or pink stain indicated
viable seed. This method has probably not been tested sufficiently for
recommendation at the time of writing.
OTHER SL-\lNLT\':G METHODS

Orher biochemical methods for determining viability have included

boiling in water with a drop of cOncentrated sodium hydroxide for from
twenty to thirt), minute... (Windisch-according to Hiltner, J891);
potassium permanganate reduction (Hibbard & Street, 1929); treatment
with weak hydrochloric acid solution (Mazkov, 1936); treatment wirh a
saturated alcoholic solution of guaiacol (Briichcr, J948); and treatment
with solutions of phosphoric acid, benzidine, or hydrogen peroxide-ill
each case containing sodium chloride and followed by treatment with
Skellysolve B (Gustafson, 1951),

FWORf.Sa.;\'CE l\.1ASURE!\ITh"TS

Kugler (1952) reponed rhat fluorescent substances leach out of nonviable seeds of RapJwnus satrous, Sinapis nIbil, and Lepidiu!1t smivullI, but
that no such subst:mcc can be detected in good seeds. Germ datn.:lgc
in wheat grain is positively correlated with the CA1:ent of browning as
measured by the fluorescence of aqucous c:\.1:racts (Sorger-Domenigg
et nl., 1955).

I20

XIII
METHODS OF TESTING FOR "lABILITY (contd.)
TETRAZOLIUM SALTS

KUHN & Jerchel (19+1) ,vere the first to recognize the value ofterrazolium
salts as excellent indicators of reduction in biological materials. Seeds,
leaves, and roots of Lepidium sativum, fermenting yeast, and lactic acid
bacteria, all became ted after treatment 'with the colourless tetrazolium.
This chemical has an advantage over dinitrobenzol, which produces an
cvanescent colour in the tissues and which is poisonous, and ovcr sodium
selenite which, though producing a permanent colour, is also poisonous.
Further, tetrazolium salts ate more stable than some of the dye.,> and work
evcn under aerobic conditions. In addition to impaning a permanent
colour, the tetrazolium salt has a low toxicity both for the plants and for
the experimenter, and is oue of the few organic compounds that is coloured
in the reduced state. In the presence of viable tissue, the almost colourless
tetrazolium salt is transformed by enzymes into the insoluble brjght~red
triphenyl formazan.
CEREJli.S ~o\."'D GRAINS

Lakon, who had introduced the selenium method of testing seed

viability, was quick to see the advantages of the new tetrazolium method,
which he developed. A description of the Lakon method for small cereals
and mai:.w is to be found in his article on 'The topographical tetrazoJium
method for determining the germinating capacity of seeds' (Lakon, 1949).
These rules for the use of tetrazolium were sllIlllIUrized in order to correct
errors in the method as used by many people after a detailed description
was published by Lakon in 1942 (Lakon, 1942, 19420.). The use of the
method requires the preparation of the embryo of the seed, so that all
those parts which are stained by the chemical, i.e. the living tissues, will
be visible. E.xcept in the case of oats, the pericarp of cereal grains is
impermeable and opaque, and the embryo must therefore be removed with
a sharp scalpel or needle before being sectioned longitudinally. In the
case of maize seeds, a longitudinal section through the centre of the
embryo is insufficient, because the lateral root primordia remain invisible.
Pre-soaking of the seeds in water facilitates the removal of the embryo.
For small cereals and maize grain a 1 per ceut solution of 2,3,5triphenyltetrazolium chloride is used. This reagent is light-sensitive, and

'"

SF..EV PRESERVATION AND LONGEVITY

so should not be exposed to direct sunlight or to prolonged diffuse light,

though solutions of it will remain unchanged for several months in the
dark. The pH value of the solution should be between six: and seven for
the best results. Embryos Cxtr:leted for testing must be free from starchy
endosperm and pericarp fragments, and must be continuously submerged
in the solution. The re3etion is complete after from seven to eight hours
in the dark at room temper::t.ture, following which the response can be
accurately determined according to LakoD (194-9). To be classed as \';ablc,
embrfos must have the plumule ::md adjacent tissue-bearing root primordia
stained. Figure 12 shows the aspect of staining for seeds of wheat, rye,

Fla. 1~.-J::xcised cmbr}"OIS of ccrc:>.Is. (,g) Wheal; VJ r)"e; (r) barley; (J) O:U$. Those p:uu
of 1~ tmbl")"o whkh, at lhe 1=. mll$l;:d>ow colGration after lrealmlml "ilb lcuuolium in
order for the kell"lello be germill:l.b1"" are sbo..n by ho:lch m:u:b (frmn.L.al;o.."'l, 1tWJ. CounCS'o'

of .'\gricuhur.ll CoUege, Stuttpn-lIoh=hcim, Gcrman~l

barle~', :md 03ts. Mai7.c grains differ in that germinable ones are stained
cither completely or at least in the region of the shoot, the smincd parts
including the initials of secondary radides and the scutdlum. Unlike the
situation \\;th other cereals, the staining of the scutellum of the maize seed
is signifiC:lnt and at least half of its area must be stained to denote viability.
Also, the tissue connecting the embryo and scutellum should be stained
in its entirety. Lakon has pictured the minimum stained arca required for
a test of germinability of maj7.c seeds in Fig. r3. Lakon warns against an
extension of the test beyond twenty-four hours, for bacterial growth rna)'
appear and be stained by the tctrazolium and hence cloud the culture. In
such cases the seed parts may appear to be stained improperly, being
covered with minute red spots. Recognition of this condition is of prime
importance, but the correct test of the seed tissue can still be determined
by removing these Spots by gentle abrasion of the surface of the tissue
underlying: them.
Many workers have been concerned with the staining of grain seeds,
because of the necessity of carly gennination-appraisal aftcr harvest and
of the tendenc)' shown by many of these forms to develop a dormancy
during maturation. This dormancy is brokcn naturally by a period of dry
122

l\1THODS OF

TfSfI~G

FOR V1ABILITY

storage, but requires from one to several months before germill:J.tion can
be obtained speedily under ord.in:J.ry germination conditions. Cottrell
(19+7) tested cereal seeds by the tetr:lZolium method and was able to
evaluate their germination capacity, either as a final or as a screening test,
within twenty-four hours-as compared with ten or more days for a
regular germination test. However, he found the method more exacting
and time-consuming for the worker th~n ordinary germination testing.
.::: ..
. .:

....
l'r

."

\~ - '..

........ -

...... _ .
. ...... '.

'-

'.

So

,.

FIG. 13.-Me<.!iau longitudinal section through a kemd of Maize. Sc, scutellum; Pr,
procambial str.md; C, oolroptile; PI, plumule; B, base of plumule; A, trlIlIsition from plumule
to ndide; R, radicle; SW, initi:lls of secondary ndieh:s. That portion of the crnbr)"o which,
:1f the It""'lSt. must ~how coloration after treatmern with tetrazolium in order for the kernel to
be gcnninablc, is shaded by hatch m:lrl.:s (from Lalo,,> 19-19. Courtc.~) of Agricultural College,
Smttg:rrt-Hohcnhcim, Germany).

The scope of the test is limited by the size of the seeds (Cottrell, 1948).
New seeds of barley, oats, and whear, incubated in a 1 per cent solution
of 2,3,s-triphenyltetrazolium chloride at 4S"C. for five hours, gave a good
colour-reaction which appeared to be a quick, reliable index of germinabilit)" but for old seeds of less than 60 per cent germination capacity, the
test was less accurate, according to Shue! (1948). Favilli (1950) used
2,3,s-triphenyltetrazoliuJl1 bromide in addition to the chloride to examine
corn, wheat, rye, kidney bean, and lupine seeds, reporting an indication of
viability which was 3 per cent greater than that secured in direct germination tests. He lists 119 items of literature on the subject
123

SEED PRESERVATION AI\'D LONGEVITY

There was an average difference, between sand germination and vigour

as indicated by tctrazolillm staining, of 7 pcr cent in tests made on wheat,
barlcy, and rye seeds (Svenson, 1952) having gennination capacities of
from 80 to 100 per cent. Sceds showing intermediate colours described as
pale pink, pink, milky red, or bluish, were interpreted as incapable of
germination, as the terrazolium indications were here usually higher than
the germination tests. This emphasize.... the importance, and difficulty, of
interpretation in many cascs. Volhen the germination capacity of the seeds
was less than 80 per cent with the sand method, the differences between
the two methods was increased, resulting in discrepancies offrom 30 to 40
per cent when the germinating capacities dropped to 30 or 40 per cent
(Svenson, 1952). Toniolo & Beni (1954-) also noted that the tetrazolium
tests were good for seeds of high quality only. The method has been used
on grains by many other people, among whom are Fink & Schweiger
(1950), Bernal (1953-54), Rirvanen (1953), and Marras (I9SS).
Rice also has a period of dormancy following harvest, and the triphenyltetrazolium bromide tcsts have been made on these dormant seeds as well
as on rice seeds which have lost their germination capacity (Niles, 1954-0
1955) It is possible in this way to determine the eventual germinability of
dormant seeds immediately after harvest, and so rice unsuitable for seed
purposes can be diverted to other uses without the necessity of storage for
periods of up ~o twelve weeks or even longer, and of periodic testing. The
tetrazolium staining provided an accurate estimate of \'iability for the
whole period extending from harvest time, through dormancy, ::md to the
complete loss of viability (Niles, 1955)'
Hyde (1949) has made a thorough study of the precision of the
tetrazolium method in the estimation of germination capacity of seeds of
Qlewing's fescue. Using a special method ofstaining and of examining and
classifying the embryos according to stain distribution, he obtained a high
correlation with the percentage of viable seeds as indicated by a germination test. The standard error of estimated germin:ltion percentage from
staining percentage was 5 per cent. When the tests were based on 300
seeds and the germination capacity was approximately 50 per cent,
differences of more than 8'5 per cent were significant. Hyde concludes
that the method is useful for seeds of Chewing's fescue and ryeg-taSs
(Hyde, 1949, 1952, I95S). In the case of .r}'C'JtaSs, there was close agreement between staining and germination tests for samples ranging in
viability from 0 to 97 per cent, and the staining method proved more
reliable when testing the secus during the after-ripening period than at
other times.
Lakon has been among the foremost proponents of the use of tetrazolium for testing seed viability, and has published many articles on the
subject (e.g. Lakon, I9So, I954; Lakon & Bulat, 1955)' The method was
received with enthusiasm by many workers confronted with the necessity
of producing quick viability tests. The results of their experinlCnts have

"4

J"tETHODS OF TESrING

ro]{

\'I,\Rl l.ITY

produced various judgements of the value of the tc;sts, from unqualified

recommendation to complete rejection.
Tetrazolium staining \,as used successfully on the seeds of ten
different kinds of grasses by Wharton (1955), and Davies & \Vinstan!ey
(1957) found that it closely approximated the standard germination tests
of seeds of Mexican grass (I.ropllorus ulIisetus).
As was pointed out by Lakon, some of the failures may have been due
to improper procedures, and variability in effectivenes,s was to be expected
as the method was extended to many other types of seeds than grains.
Also, as other seeds arc used, special adaptations of the method must be
made to fit the characteristics of the seed under study.
There has been a great deal of interest in the use of the terrazolium
staining method for detcrmining freezing injury to maize seed, a subjcrt
of some economic importance. The method proved a reli:lble index to
germination in the testing of four samples of artificially frozen, and one
sample of naturally frozen, hybrid seed maize grain. These grains had
been frozen at 48 per cent or less moisture and then dried to 12 per cent
moisrure before testing (Goodsell, 1948). The staining followed the use of
025 per cent tctTaZolium solution to which was added 1 ml. of IO per cent
sodium hydroxide solution per IOO ml. The minimal damage of a hard
freeze to viabilit), of the seeds in a hybrid seed maize field could be
determined within forty-eight hours if the seed ears werc carefully
shelled by hand and the seed dried rapidly to 12 per cent moisturc with
heated, circulated air. Isely et aJ. (IlJ.j.8) harvested maize seeds at various
stages of maturity, subjected them to various sub-freC",6ng tcrnperatures,
and tested them with tetl:lZoliurn chloride using standard germination in
sand as a control. They found that tetrazolium gave higher readings in most
cases, e,,>pecially with high-moisture seed, than did standard gelmination
in sand. Also, interpretation was difficult with frosted seeds and had to be
made on the basis of differential staining nther than on the presence or
absence of staining:. These authors concluded, however, that the substitution of staining for germination tests is justifiable when immediate
inforIrultion is necessary.
Bennett & Loomis (19-l9) found the tetrazolium tests to be fairly
accurate in assessing freezing injury to maize grain only if the germination
was moderately high and the grain had been stored for some time after
freezing. Necessity for storage may be related to the moisture content of
the grain, which was 30 to 60 per cent in freshly frozen seeds, and which
affects the staining test (see Goodsell, 1948). Bennett & Loomis (1949)
point out, however, that it is possible to estimate the proportion of injured
seeds immediately after freezing. This is done by observing the tint,
texture, structural appearance, and uther characteristics of the injured
tissue-instead of the presence or absence of a pink stain-but this
requires experience before a reasonable estimate of frost damage can be
made.
12 5

(a) Typical seedlings of Hlaek Valentine I3cans from favourable (lrfl) and unfa,ournble
(righl) storJge len days after emergence. (From E. H. Toole, Toole & Borthwid.:, 1957.)

Pun 10
(b) TypiClI plan15 of Black Valentine lkans from favournble (lrft) and unfa,ourable
(right) stonge at m:uuritr. (From E. H. Toole, Toole & Bonhwiek, 1957.)

Ph(/{os by lQurlrq Unitrd Stairs Dlparlmrnl oj AgrirofllUr

~lTHODS

OF TFSfING FOR VI,\BlLITY

TCLraZolium c:hloride also offers a rapid means of testing the "iability

of cottQIl-sct:US, though ;lll)' rL':lCUOn Other than complete staining of thc
embr)'o is indicative of loss of viability (Lambou, 1953).

TREE AND SIlRun SEEDS

It woulu be an advantage if dormant trce-seeds could be tcsted quickly

by some less tedious method than the excised embryo technique.
Nauvornik reported as early as 1946 that tctrazolium staining tests were
fe:lSible for Lhe determination ofyubility ofseeds of fruit trees and shrubs,
good agreement being obtained between such tests and regular germination tests (Nadvornik, 1946).
An extensive comparison of the testing of both dormant and nondormant seeds of varying viabilities of woody plants by the cxcised embI) 0
and tetrazolium methods was descrihed by Flemion & IJoolc (1948). They
used 100 different lots of seeds representing fifty-eight species and
seventeen families of phnts. Detailed results of these experiments arc
sho",TI in Tables :A'V, XVI, and ~\'lI, wen from theartide by Flemion &
Poole (1948). The 'rapid ,;abili~' test' referred to in these rabies is the
excised embI)'o technique. It will be noted that the tetrazolium tests ,",cre
onen characterized by different degrees of staining as well as by spot
staining of all or some portions of the ,-iable embryos. Dead embryos
rarely became stained. The excised embryo tests also indicated two types
of germination response, IUmely vigorous and sluggish. The data have
becn combined and rearranged to show general effects (Table X\I11I).
The amhors c\':plain this table as follows (Flcmion & Poole, 1948, p. 25 1):
'The number of samples (column I) \\-ith a particular type of staining
(column 3) which showed various degrees ofyiabili[}' (column 2) by the
rapid viability method are recorded. These arc compared with the percentage of staining obtained. In 9 to 19 per cent of the 100 samples
the \iability test and the staining test agreed exactly (columns 4 and 5.
lines 13,9,5. and I. from the bottom). In a third to about a half of the
tests thc tetrazolium readings were from I to 30 per cent above or below
the percentage viable obtained by tbe rapid viabilitr test. In a considcrable
number of the tcsts the deviation between the two methods was grcater
than 30 pcr cent. For instance as in the case of embryos completely sraincd
including all shades [Table XVlIIJ (line 9 from the bottom, columns 8 and
TO), one-half of the JOO samples tested deviated more than 30 per ccnt
abovc or below the rapid viability readings.' The corrclation between the
two tests is significant when all the data are combined in this way, hut the
variations in individual tetrazolium lests ma.de some of them of doubtful
,"alue. The conclusion from these comprehensive tests is that more work
would need to be done on an individual species before the staining test
could be used as a measure of germination capacity.
J27

TABLE

XV

RESULTS OBTAll\'ED WITH RAPID VIABILITY TEST AND STAlNING WITH 2,3,5-TR1PHF.NYLTETRAZOLIUM CHLOIUDE IN DETER.\llNI!\G

1HE VIABILITY Of VA1UOUS SEEDS

TCllm'.oHum test
giving colour

l'~r ~e~1 ofsccd~

Rapid viubility te,l

Family

Accf:lccac
Anncnrdiaccae
IJcrhcridaecac

"N

00

Bigll0n;accac

Arer Jar/nri,l/III L.
Rhus uramil/irll Air.
1J~rh{ri$ tllllllhergii DC.
lJ~rhtris l/mnhrrgii DC.
Berberis illllnbcl'gii DC.
Campsis radiams Seem.
Calalpa spcrios/J Wuclcr

Caprifoliaeeac
O:lastnlccac

l,fJn;am diojrQ L.

Cornncc'lc

Cornus sp.
lJolllamriis lIirgilJilm/l L.
IlllllJamriis t'irgiIJilJ71a L.
Lilfllidamb/lr s/yradj/u(I L.

I-lamamclidu~-eae

Le,uminosuc

C~faSir/l$

RQhi,lia pscudo-acacia L.

M\\lvaccac

9'

, 90
, ,,,

,,,

Tot.1!

6
4
4

gil
,, , S4,
", 7'"
7'
, , ,
77

9'
S'

s(alldms L.

errds chillemis L.
RQbinia pstudlHl~acia L.

Entire embryo
vigor- slug- %
sminell
O\IS gish liable Deep Med. Light

Species

73

,,
,

Abrlmoschus mu/mlus Moench (Okro)

GOlSypium IlirJulJilIl L. ~Cultun)
CoslJpilll1/ ilirmllllll L. O)[(unj
GossJ'Pitill/ ilirSUlrml L. (Omun
fJibjs~us syriatlll L.

'"
II
6;
9'

77

8,

76

34
60

It

",

"

1,

, ",
63
'7,
96

49

"

"

48

55

54
9S

""9

3'

,6

""

37
H

83

95

So
,8

"'3

,,,

'7

"

36

,,

~"

,6

<4

'9

'4

,6

9
5

7
7

"

'4

,,

J'

'00

'"
'00

,
8

'00
'00

4'

"

95

"

4'

"

'9

74

43

J7

'00

7'
OJ
8,

nteJ or
Most of embryo
Spot
No
empty
smincd
stnining
stain- secds, tlot
D~cp iVied. Light Deep Mcd.
Light ing ["Mod, %

95
54

DClcrior-

,8

, "
,
,
i~

,
,,,
,,
,
0
0
0

!,

,8

~
<

>

:i
C

~
~
~
~

METHODS OF TESfING FOR VIABILITY

0000""""'0

<'.

"

"

" .-

"

,
M

" "

"

,"

o
-e,0

00

,
'"

o
I

TABLE XV!
lU::SULTS OIlTAIl\"ED WITH GERMINATION TESTS, RAPID VIABILITY 1'8f, AND STAU,ING WITH

2,3,S-TRIPHENYLTEl'ltAZOUUM

CHLORIDE, IN DETElUIIlNING THE VI,\BILlTY OF VARIOUS PINACEOVS SEWS

Germi-

Species

Halion
[cst -

,X

vIgor-

sluggish

OllS

Abi(J bi/!s//I/I(I/ Mil.

LaTi,r turopaea DC.

Pirm c.tulsa Link
l'irm (xu!sa Link
.... 1';(((1 rx(~IsII Link
/'irru gl'lllc(l (P. ClJlllldclIlisllS, r.
$lmul/)
l'ICt'll pur/grns Eng-elm.
/';UIl plmga/f Ellge!m,
Piera pi/Ilgens Enlj:elm.
Pinus ballf.:Jiulul Lamb.
Pim,s tprjfl(ltQ Morelet
J'iw/s mribpfQ Morelet
PirmJ cu,ibllfO Morelee
Pi,HlS cu,ibilfll Morelel
Pimu crhinala Mill.
PimlS crhinala Mil!.
Pimis Plllusl,is Mill.
.PiIIllS 5Irab"J L.
PitlJlS .lylvesl,is L.
Pitltls .ylves/,is L.

15

Pilllls syfVtJ/,is L.
Pillus IIJ.cda L.
PitlllS Ilmia L.

\':11'.

-"
-7'
"l'

13

-6,
--7'

--.6
-

Tetr"zoIiulIl lcst
PCI' CCnl of seeds giving colour

Rapid viability test

"",
,

7J

3'
1
7J

7'
39
6-

"
gg

",8
4'

7'

t:ii

..

,6

,
,
,,
,
,,
,5
,,
,,
,,
",,
.1
,,
0

Total

viahlc
34

,8

,,

73

Entire embryo
Most of embryo
SP'?t
No
suiocd
Stained
staming
st:ljn~
D~'Cp r,.Icd. Light Deep Mell. Light Deep Med. Light ing

37
,6
53

'5

3
1

, 'I ,
83

38

"

7'
39

6,
,8

"
61

,6

86

~I

"" ,8
i1, "
g

77
47

7'
3'
'3

"

"

"

'3

'7

'3

3'

"
3'

3
3

",6
"
u

"
"

itiliilil:biUiiriiiHllliiUwii"Uhii"iil~t"Ui'~llwilllWiiiU*UiiHi!il~ri;h
j;j;!U*1ji ih;;;;ii;+ittllUt;,;Ul,.I'jiUjil !liiIMj

"

74
3

, I;,
,

6
3

34
33

,
,6

,6

",
6

"3'

,8

IC!i[cd,

,8

3;

61

seeds, nOt

,
,,

"

57

Dctcriomtcd
or empty

"

"6
8
""
l
"'3

'.
'. .'

"

"5

'7

.8

,~

~~

~
>
Z

0
r'

~
~

METHODS OF TISrING FOR VLWILlIT

,,~

'0

'" r- '':; ~"

.,.,
"" "
'''~
~

,,~

;;..,:;

,~

""

."lOtl 00

I I

00

I I

'3'

TABLE XVII
RESULTS OIlTAII\"'ED WITH RAPlD VIABILITY TEST Al\lJ) STAINING WITH 2,3,5-1RIPHENYLTETRAZOLIUM CHLORIDE IN DETERMIKI~G
THE VIABILITY OIl VARIOUS ROSAaoUS SEEDS
Tctrazolium lcst

Rapid Villbility ICst

Species

,X

TOlal

Ylgor-

slug_

ous

gish

viable

J"

",

+I

C)'J,mill oblollga Mill. (Quince)

3'

Malil. canlllltlnis DC.

MalliS jloriblmda Sieb.
tIIalus J{"dd~ck(fj Spaeth

0
0
0

"

,
CllurllQmeles japonirQ Lind!.
Clri/mollltles japonirll Lindt.

(Jnpan-Q!Jjnce~
(Japan~incc

... Prlnsepill sitUllsis Olivo

w
N Prl/lltu

1/III(ricalla

M'lrsh

(PJuml

I'rlll1ll1 amrricallil Mnrsh (Plum

Primm um~rirulla M.arsh (Plum)
I'rulIIll aviulII L. (Ma~~a~
I"IIIIIIS allium L. (Ma~~ard
l'rmllls'trasiftra Ehrh.~, yrobolllll Plum)
l'rtlUIiS &trasifira Ehrh. Myrobotln Plum)
I'rlmUJ muftllltb L.
Pr/l1tt1J malta/"b L.
Prum/j pffJica (L.) Stokes (l'caCh~
PmmlS pffsi&a (L. Stokes (PCllch
l'rtmus prrSi&fl
Stokes {Peach
PrIllWS I'irgiuiartll L. (Chokecherry)

(L.~

Pyrus sp. (Kiefli,r I'ear)

L. (E"",,,"

'0
0
08
97
0

",."'hl

,"wp,,;, L. (European i\11..ash

SQrb/1I flu(l/parifl
SUr/illS auwpari" 1.. (Iiuropean l\1t.-ash
Sorblls aucuparja L. (European J\.lt.-ash

3J
3'

0
'3
0

"3'
30

6
0

"

36
,8

"
79

""
8,

"

70
;
7
'3
0

"
0

,,

6;

'3

'"

5'

"

7"

,6

"
'00

,6

" "
"

"

"

"

'"
5

,6

'9

'5

98

"

'9

'5

6,

'"

'"

45

"

!J<'

"

'00
'00

"

3~

60

"

"

57

"

"

79

1\2

,6

7"
'3
0

67

'00

"

"

"
S~

", ,

"

sccd~. llot

tested,

",

"

,
,

0
0

!I.:,illlllll,llllllllllllliilll

~
~

>

S
Z

0
0
0
0
0
0
0

"

'00

63

>

0
0

37

......liliU.i..l.lliilhu......................... , ..................H........................... II ..UIUI .. UUUIUIUUUiUU.UU.uiiUWUi.

Dctcrioutcd
or empty

'00

'00

"

9'

N"

stained
stauung
sfdinDeep Mod. Light Deep JI,'1ed. Light Deep Med. Light ing

"
9'

0
87
63
0

Sr~t

.lI,lost of embryo
swilled

6;

,8

36

Entire embryo

3J

Rosa lIIu/lijforlJ Thunb.

S"'"

"

"
"

I'er celli of seeds giving colour

0
J
0
0

,\l!."'THOD's

m'

Tt.srtr-.G I-OR VLWILITY

TABLE XVIII
COI'o.1PARl,SON 01" 1t....<;UI.TS

BETWE.f..~

RAPID VIAfHLlTY AND TETRAZOLIUj\'l

Tt:Sl'S

No.
of
~~

pl~

.R:!pid
\iability
tcs[, %
\'l:Ibk'"

Tell"ll7,olium tests
P=nl.:lge difference betwetn
results of rnpid viability ami
{etrn.olium tClots
Sroining
sronuard

0%- 1-30% 3 1"""70%

(results
identical)
No.

45

"

,6

(,1-100
2I- 60
0- 20

Elltire embC)o st~incd

deep :md medium
rolollT

45
'9

,6

61-100
2I- 60
0- 20

Entire emhryo
mained, all
shades

3'
22

'"

61-IOOt
60t
0- 20t

;<.}-

Entire embC)-o
stained, all
shades

",6'9

6'-'00
2I- 60
0- 20

Entire embryo or
most of embryo
stained, all ..hades

"

%
'5

38

"

"

"

''".

33

33 '7 '7

" "

3'

,.

9 35

" "
9

.)

""

54

0'7134

,8

" "

87 5 '3
" " 33
,8 " "
n
,
" " , -", , , '97 3'
'9
48

"

,
,

'"

3 37 8,
7 7 '4
38
4

'4 '4 45 45

'00

9 '0 H
'0 35 n

'9 '9 37 37

'9 '9 '3 45

'00

No.

33

'00

"

70

33

'00

% No. % No, %

, , SJ
, 3 "7 g
,
9

>

Coefficient
of
corrd.ation

" '"
5

3
n

o'86Jo

,6 ,6

' ,-

3'
4 '4 54

0-7<).;<)

08202

30 3"

'" All of the embryos showing any dewlopment were listed as viable.
t

Only embryOS sbowing \igorous development were considered

:IS

viable.

Since 1948, the use of 2,3,s-triphenyltetrazolium chloride as a useful

viability indicator has been reported for seeds of conifers (parker, 1953;
Yanagisawa & Asakawa, 1953), Beech (parker, '953; Evrard, 1955), and
oaks (Parker, 1953). Hilf & Rohmeder (1955) give a derailed description of
the penetration by diffusion of terrazolium into the different cells of the
embryo and endosperm tissues of forest seeds, and of the rapid reduction
of the chemical in all tissues.
1I-IECHAi\,11SM OF AGTION OF TETRAZOLIUM

Tctrazolium salt is an oxidation-reduction indicator, and the development of the non-diffusible red colour in a specific tissue is an indication of
the presence of activc respiratory processes in which h)'drogcn radicals
are traIL"ferrcd to the terrazolium chloride. The reduction of tetrazolium
to the coloured formazan by living tissues is the result of enzyme action.
e. O. Jensen et ai. (1951) thought it probable that the reduction of
133

SEt:D PRESt:RVATION At'\"D LONGEVITY

triphenyltetrazolium chloride by \;able maize embryos depended upon

onc or more dehydro<;enase systems with favourable redox potentials. As
we have seen above, the presence of active enZ)'IDe systcms does not
necessarily indicate Yiabilit)', but thc absence of dehydrogenase activity
indicales inability to germinate. Roberts (i95I) agrees ",ith this explanation of the staining effect. The reduction of terrazolium chloride in barley
seeds is catalrsed by a dehydrogenase, or debydrogcnases, with pH optima
at 7.0 and 7'7 (Macleod, 19Sza). Although the loss of dehydrogenase
activity in barley seeds paralleled the loss of ~ability, germination figures
were apt to be lower than the yalues of a.ctive dehydrogenase indieated.
Furthermore, enzyme acti\'ity persisted in hcat-damaged barley seeds
after complete loss of germinative ability_ l\'IacLcod concluded as a result
of these tests that this staining technique is not a measure of life in the
seeds but rather of the presence of active organic acid dehydrogenases.
F. G. Smith (1952) made a detailed study of the phenomenon and
concluded that a positive tetrazolium reaction in the secd tissues may be
due to the action of pyridine-nucleotide-linked dehydrogenases or
possibly to aerobic dehydrogenases such as xanthine oxidase. A weak or
negative reaction, Smith says, may indicate a deterioration or lack of the
dehydrogenase or of diaphonse, which is thought to mediate (control)
the reduction. An insuffici~nt supply of substrates, or competitive action
of a~robic hydrogen acceptors, rna)' :llso result in a weak terrazolium
reaction. Under conditions where rerrazolium is used as a hydrogen
acceptor, it may cause significant alterations in the respiratory mernbolism
(Throneberry & Smith, 1953).
Ingcncral, itan besaid that staining with terrazolium has an important
role to play in rapid viabilily tests, especially- in the cases of seeds of
cereals, grasses, vegetables and other economic herbaceous plants; bur
the best method must be developed for each variet)', and interpretation
of results is much safer in the hands of experienccd technicians than of
others. A recent description of the general technique of tetrazolium testing
has been given hy Moore & Smith (1957). Fink & Schweiger (1950) give
a table of results of tetrazolium tests by other workers. It has been generally
conceded by workers in the field (see- above citations; also Parker, 1955)
that, although reduction of triphenyltetrazolium salts is nOt always a
reliable indicator of the percentage germination to be e."pected, staining
failure in cells normally reducing thc chemical is a certain sign of
deterioration. Whether it can be used as a general test for life in all
tissues is still a qucstion (Manson et aI., 1947).
That the tctrazolium method of testing "iability has great potential
importance is secn by the appointment of a Biochemical and Seedling
Vigour Test Committee by the International Seed Testing Association.
The 1953 report of this committee (Gadd, 1953) gave the results of
experiments b~' seven different stations on seventeen samples of cereal
seeds and one sample each of pea, bean, pine, spruce, and pear seeds. It
134

METHODS OF TESTIl'\G l'()R VIABILITY

was found that the test could be carried our with precision at the different
stations and that, in general, similar results were obtained at these
stations. The staining test did not reveal serious damage to rye grain or
give information about low seeding value, but was a good index of sowing
value of high-germinating seeds, and of one lot of medium germinating
capacity. As a result of these co-operative tests, the committee concluded
that they could not recommend the adoption of the method to replace the
germination tests of cereal grains, though it could be used with success
with other seeds. In fact, the use of the terrazolium method for certain
kinds of seeds was included in the Rules agreed upon by the International
Seed Testing Association in J953. These rules made it obligatory to
include tetrazolium tests for viability of Carpinus, Pinus cembra, Prunus,
Rosa, Taxus, and TiNa, and permitted their use for Fraxinus and}uniperus.
The r957 report of the Biochemical Viability Test Committee
(Germ, 1957) includes results of ten comparative tests on these seeds. The
data from these tests and information from a questionnaire sent to seedtesting stations led to the following 'Synopsis and Suggestions' (Germ,
1957, pp. 320- 1).
Biocllemical tests are already made by a considerable number of sCt':d-tesling
srations.
z. Amongst chemicals used for this purpose, tctrazoliuru-sa.lts are taking fin;l
pillee.
J. The topogrnphy of the colouring is decisive for the judgment.
Uniform results can be acmc\'oo only if standardized meLhods are UStd.
Special methods must be worked out for every kind of seed.
5. It is ret:ommended 10 include the methodology worked out by Dr. Bulat in
its rules and make it obligatory.
6. There exists no reason why the tetr:lZolium-method should be remored from
the rules for those kinds of seeds for which it is now obligatory or pcnnissihle,
as long as the method to be used for each kind is well defined in the rules.
7. It is recommended to permit the tetT:l7.oliuru-test or make it obligatory in
future only if the method for that particular kind has been exactly worked uut.
8. The analysts who are carrying out the tetr:lZoliuru-test must be adequately
trained.
9. It is recommended to draw attention to the difficulties of the tetrazolium-test
either by the following sentence or some other senlence lo the similar effect:
'The tetrazolium-tcst should be carried out exclusively by experienced and
sufficiently trained analysts.'
TO. It should be suggested to the scro-trade (FIS) not to finalize purcha,~~ based
on tetrawlium-\-alucs, and to make reference to:tn arbitrary analysis if possible.
11. The latitudes ofanalyses as tbey arc included in tlle rules for germil1:1tioD-lests
can be used for the tetrazolium-tt.'STs only if these arc carried out wilh 4 X roo
seeds. It is recommended not to depart from this rule in the tetra.zolium-fcsts.
J2. We have seen that it is possible to bring about germination of kinds of seeus
which can otherwise be genninatw only with great difficulty by certain types
of preparation (Tilia and Frarimll). Further efforts in this line should be made
as they might lead to the disco\'er)' of some simpler and eheaper method than
the tetrazolium-test of testing the vitality of difficult seeds.
r3. The further aeti\ities of a similar Comminee in this line should include detailed studies of the methodology applicable to those types of seeds which are
I.

'35

SEED

PRESER\'.\TIO~

Al'.'D UJ",GEVITY

already included in the rules and new ones to be included, and the initiation
of an enquiry amongst aU interested members.
WCfR1C.U.

R1::sPoNSrS

Wallcr (1901) attempted to estimate the vitality of seeds by means of

an electrical 'aftcr-currcnt' excited by living matter as the result of an
'exciting current'. Fresh, vigorolls bean seeds gave a large response, while
dead seeds or those of low quality gave little or no response. Similarly,
good seeds killed br boiling produced no currents. In general the 'bb.ze'
reaction of the seeds was parallel to the results of germination tests. Also,
the correlation between the electrical reaction and the germinative
capacity led to the conclusion that the magnitude of tht: reaction is some
measure of vitality or vigour of gennination. Fraser (1916), using the
same method with seeds of barley, confirmed the results of Waller.
Fick & Hibbard (1925) adopted the mcthod of measuring the rehtive
outward diffusion of electrolytes (as indicated by conductivity readin!,rs)
as an index of viability of timothy and red clover seeds. They considered
this method superior to the mC3SUIement of resistance of the seeds them~
selves and to the comparison of the relative absorption and excretion of
salts. This method was examined further (Hibbard & Miller, 1928) and
found effective for disrm.,"l1ishing bern-een seeds of high, low, and medium
germinative capacity. However, these authors noted objections to the
method. It is expensive and requires an operator with general knowledge
of conductance measurements. It also presupposes that the electrolytic
contents of aU samples of one species are the same, which is doubtful, for
the chemical compositions of seeds are lmOl'o'Il fa \-ary according to the
locality in which they are gro",'ll. The chemical composition of seeds is
especially sensitive to nutrition, which varies greatly in different soils.
The 'prime' potential, or the reading obtained when the electrodes
first touch the maize seed, is highly correlated with seed viability as
mea.<;ured by percentage germination and by yield (0. E. Nelson & Burr,
1946). The 'equilibrium' potential, which is reached after the galvanometcr needle has fallen off rapidly for from 30 to 120 seconds, and which
is rerained for twO to five minutes, is nOt related to seed Yiability, but
rather to the generic constitution of the plant. Seeds with superior gro"--rhperformance could be segrC"o'ated rt";uhrly by this method, evCJ1 though
such growth was not manifested by the plant for from six to eight weeks.
The electro-motive force of the seedling can be used to determine the
inherent vigour of various maize hybrids :md hence could be of great
wue in genetical work, as well as in determining the best prospects for
survival in the nursery and garden (Anon., 19+7).

X-RAy

A."l..u.YSIS

Pcrhaps the most recent of the rapid viability tests is the X-ra~T
contrast method developed by Simal: and his co-workers (Miiller-Olsen

136

METHODS OF TESTING J'OR VHDlLITY

et a/., 1956; Simak, I957; SiIDakell1/., 1957). It has been used exclusively
so far for conifer and other tree seeds (see above citations; also Evrard,
1957) The method is based on the penetration of the seed-eoat and of the
dead seed-tissue by barium chloride In hydrous solutio~ followed by a
radiograph which reveals the necrotic, impregnated parts of the seeds in
striking contrast to the living, unimprcgnated parts. Various impregnation
patterns of seeds of Scots pine (Pinus sylvestris) are shown in Plate 9
(Simak, 1957). Simak point,> out that photographic enlargements of
radiographs are difficult to reproduce and hence sketches arc included in
the figure.
The X-ray method has an advantage over other quick vjabili!)' methods
which require the removal of the seed-coats, and, in many cases, some
special preparation of the embryo for testing. Also, the X-ray contrast
method offers a combined analysis of the structural and physiological
condition of the seed. The former is asccrsained by the relation of embryo
and endosperm shown in the radiograph. fu the seeds werc not killed by
the X-rays, it was possible to lay them out in the germirunor in exactly the
order_shO'\ll in the radiograph, and thus consider the germination of
every single seed on the basis of its anatomical strllcture and the degree
of impregnation. Sinuk (1957) found a single seed of onc sample of Sco!.S
pine which genninated in spite of its impregnated endosperm and
cotyledons, thus showing the test to be not infallible, and he suggests that
the X-my test may be combined with the tetrazolium test. The two arc
complementary in their principles and thus a tOpo&'Taphic classification
for the X-ray method could be evolved. An impregnation ratio of onefourth has been defined to mark the border between viable and nonviable seeds. Simak (1957) warns that barium chloride is not a universal
impregnation agent for X-ray contrast~nalysis, and that seeds of different
species m:lY react differently to the same chemical substance.

'37

XlV
PLANTS FROM OLD SEEDS
THE increasing practice among sccdsmen of keeping surplus supplies of
seeds for sale in subsequent years has resulted in pan from the experimental determinations of the storage conditions clfcctl\"c for the maintenance of viability of the various types of seeds. FOT the most part, the
measure of the keeping quality has been the power of the seeds to produce
seedlings as determined by germination percentages obtained from soil
or other plantings. However, the vigour of the plants produced from old
seeds is a matter of increasing importance in vicw of this mounting practice
of storing seeds for long periods.
There have been reporlS from time to time on the \"a1ue of old seeds
for planting, apart from their germination capacity_ Bartlett (1859. p. 332)
quoted from the irish FQTmu's Gauttt that, 'The gardener ImO\\S that
melon and cucumber seeds, if used of the last rear's saving, produce
plants too vigorous to produce much good fruit; whereas, those kept
over for several years produce less rambling, but very fruitful plants'.
Ohga (1926; see a/so Plate I) found that sprouts from Indian lotus
seeds reputedly more than 1,000 years old were always ]ongt:r than those
from new seeds of this plant. A note (Anon., 1932) on trials to determine the relative values of pumpkin seeds of different ages, claimed that
three-rear-old seed gave the best results in one trial while, in another,
four-year-old seeds yielded less quantity but better-quality pumpkin
fruits than one-ycar-old seeds. It is not srated whether the yield was from
a given number of seeds or from a given number of plants. Delayed
germination, followed by decreased growth-rate and lack of a wellde\c!opcd primary root, characterized old seeds of cucumber (Parkinson,
1948). Old seeds of lettuce also produce abnormal, very short seedlings
with practically no roots, according to Drake (1948).
Beattie et al. (1932) determined the yield of peanuts from old seeds.
They did not measure yield from the same number of plants in all C;L.<;es,
but rather the yield from the plantS produced from a given number of
seeds-regardless of the germination percentage. There were rather wide
differences in the stand resulting from differences in percentage of germination, but these differences did not appreciably affect the jield. Beattie
et a/. attributed these results partlr to soil heterogeneity and partly to the
larger yields of plants adjacent to missing hills, but warned that these
138

PLAl\,'TS FRO.\1 OLD Sf.EDS

results should not be taken as an indiC3.tion that the use of poor seeds is a
profitable practice.
Kicsselbach (1937) used maize seeds of various ages and tested the
yields obtained from planting them. He found that satisfactorpesults may
be expected from well-m:.ltured, viable seed maize grain up to four years
old-proYided it ha.<> been kept free from e.''{temal moisture and insect
or rodent injury. He tested seeds from one to four years of age annually
for fiye years. No significant differences were obtained in the comparative
acrc yield in bushels from seeds of various ages. }Jso, there were no
material effects of seed age upon the vegetative development of the
plant.
Dungan & Koehler (1944) found that a decrease in the yield from old
seed maize grain was caused by a reduction in the number of seedlings
established in the field, and to a lesser e.'\."tent by decreased yield
per plant. Three-year-old and one-year-old seeds of six different lots
all produced perfect field stands, i.e. the number of plants was not
affected, but the yield of the plants from the older seeds was 4.8 per cent
lower than that from the one-year-old seeds. This reduction was extended
t078 percentwhenthcreduccd stand was also allowed toinflllence the yield.
Leus (1930) studied the relation of farm-crop seeds to production and
found that in wet-season culture in the Philippines the maize ear and
mungo seed yield was greater from thirteen-month-old than from threemonth-old seed, but in the yields of seed from cowpea and soya beans the
reverse was true. In dry-season crops also the oldest maize seeds (thirty-two months old) resulted in the highest production of ears to the hill, but
cowpea, mungo, and soya bean plants from seeds eight months old gave a
higher production than those from seeds eighteen months old.
Crocioni (T934) conducted soil planting experiments with TritiClllll,
Bm-ssica, Medicago, and Tri[oliulIl pm-tense, comparing seeds of different
ages and follo\\ing the development of the plants in each case. Plants from
old seeds were retarded in growth and less resistant to adverse conditions.
Paralleling the age of the seed, he found that old wheat and Brassica plants
were smaller and their yield was decreased. With leguminous plants, the
decreased yield was less evident and became marked only in the cases of
vcry old seeds.
Riddell & Gries (1956) suggested that variations in the growth-rate of
spring wheats developing from seeds of different ages is related to the
temperature obtaining during maturation rather than to the age or
conditions of storage.
A tendency has been noted in cotton for older seed to yield less, but
these differences were not statistically significant (Christidis, 1954). Tt
was concluded that the effect of age seemed negligible for the first tcn
years of storage of these seeds. No consistent effect of age on earliness,
lint length, or ginning out-turn was demonstrated, though extcndcd
storage beyond ten years may show some such effects. Age not only

'39


SEtD

1~\'ATlOX

.-\1W LONGEVITY

decreases the viability of the embryos, but reduces the h'TOwth of pbnr:s of
tob:lcCO from ten-ycar-old seeds (Cirkovskij, 1953).
The inlluence of the age of sugar-beet seeds on their seeding ,'31uc was
studied by Filutowicz & Bejnar (1954). They conducted hotbed, greenhouse, and field experiments on eleyen nrierics from seven suceessiyc
crop years. The resultS showed that ooc- to four-year-old seeds did not
differ in viability, bur older seeds (five to seven years old) declined in
germination capability. Associated with this decline was a later appe:mncc
of seedli.n;;s in the field; bur there was no reduction in the yalu~ of beets
produced from such seed.
1JEA.t'1S

Rodrigo (1939), working "ith mungo (Phaseo/lls aI/reus Roxb.), used

eleven- to thineen-ye::J.r-old seeds which had been stored in bottles sealed
~ith paraffin w::L.'l: at room temperature. As controls he used onc-month-old
seeds taken from plantings of the old seeds made previously. At maturity
the plants were harvested and data werc takcn of the number of pods
produced, the weight of the dry pods, the weight of the straw, and the
weight of the seeds per plant. rn garden ploes he obtained a significant
difference in plam yields of new and old seeds in favour of the laneI'. Old
seeds also gave a greater pod-yield. The cffect on the yield of beans was
similar to that on the yield of pods-except in one case where the pod- and
bean-yield of new seeds \v:lS found to be slightly greater than that of old
seeds. He docs not gi"e the genninarion percentages obtained for the
various seeds used. He offers two possible explall3tions: (II) the yigour of
the seed expressed in yielding power is correlated with it.o; inherent
longevity; or (b) the seed requires some 'seasoning' or 'curing' beforc it
attains the peak of its vitality.
A recent report by E. H. Toole et al. (1957) also deals \\ith PMuolus
(three varieties of Sllilp beans), using seeds of the same age (forry-scven
months) and noting the effect on ;''l'Owth and production of seed storage
undcr favourable :md unfavour.lble conditions. The favourable storage
conditions were in a naturally dry warehouse at Filer, Idaho, and the
unfavourable stOrage was controlled :It 19"C. and 57 per cent rclatiyc
humidity at Mercedes, Texas. The seeds \\CTC germinated and grO\nl
under controlled conditions of nutrition, temperature, and light, in
randomized blocks in a growth room. Seeds from unfavourable STOrage
conditions were planted two days earlier than those from the f.wourablc
stonge. conditions, because of the difference in rate of germination and
,-igour of growth. The plants were grown until the oldc:sr pods wcre ready
for harvcst, when the entire plant was cut at ground-lcycl :md nrious
measurements of gwwth and yield 'Were made. That rhere were marked
differences in the rate and vigour ofgrowth is shown in Plate 10 (a) and (b).
Plants from the bener sced-storage conditions reached lhc flowering and
fruiting stages earlier, and gayC a significantly higher ~-ield, than those

'4

PL-\NlS FROM OLD SEEDS

from unf:lYoumble conditions-in spite of the earlier planting of the

bUeT. Thus there are differences in vigour of the plants grown frum snap
be:lll seeds which have been subjected to different storage conditions for
the same lel1h->th of time.
VARIOUS

--

OTHER

PLoI,.l'\'TS

It has been pointed out that sL"(-year-old carrot, eggplant (aubergine),

onion, tomato, and lettuce seeds (Barton, 1939(1), and ten-year-old
Delphinium seeds (Barton, 1939b), all pr{)(iuced normal plants, but no
quantitativ~ data were noted.
A carefully controlled experiment was undertaken in 1946 (Barton &
Garman, 1946) to determine whether the age and storage condition of the
seeds of China aster (Cllllisteplllis c1JillellSis Nees), verbena (Verbel1(/
teucrioides Gill. & Hook.), pepper (Capsicum [mtescells L.), tomato (Lycopers;cum rsclilentulII Mill.), and lettuce (Lactuca satiiJa L.), would have an
effect on the yield of plants grown from such seeds. In order to have fresh
seeds of the same genetic strain to compare with those which had been
storcd for longer periods, plants of the five species concerned were grown
in 1944 from seeds of the original lots which had been stored several years
before. From the..<;e plants seeds were collected, cleaned, dried, and stored
in the laborarory until April 1945. All of the fresh seeds, excepting those
of lettuce, were produced in the greenhouse, pollination being controlled.
The lettuce seeds were produced in the field hut no other lcthlce
flowered in the vicinity; hence the strain remained pure. Very fresh seeds
of pepper and tomato were secured for planting in April 1945, from plants
(also produced from old seeds of the original lots) which had been gruwn
in the greenhouse during the winter of 1944-5.
The effect of various storage conditions on the germination capacity
of aster and verbena seeds was reponed in Chapter VI (Banon, 1939b).
Those seeds '\\>lth approximately 4 per cent moisture content that had been
stored in tin cans at - 5C. were used. Plants of both of these kinds were
grown in the field in 1945 from seeds as follows: lot A, seeds produced in
the greenhouse during the summer of 1944, and lot B, seeds from the
original stored lots. Thus the seeds used to produce plants in lot A were
approximately seven months old while those used for lot B were approximately nine-and-a-half rears old.
The germination capacity of the stored seeds of pepper, tomato1 and
lettuce has also been described (see Chapter VI; Barton 1 1935, 1939)
Pepper seeds which had been mixed with calcium oxide to remove
one-half of the original moisture and !.hen sealed in tin cans and stored at
- 5C. were selected for these tests. To_produce plants for field performance tests in 1945, fresh and old seeds were used as follows: lot A, seeds
produced in the greenhouse in the spring of 19+5 from plants grown from
the seeds stored in 19321 as described above; these seeds were harvested

'4'

SF.F:D PRESERV.HION .-I..,'Hl LONGEVITY

not more than two weeks before sowing for the field test; lot E, seeds
produced in the ~'Tccnhousc in the summer of 19+4 from pl:mrs grown
from the seeds stored in 1932 as described above; these seeds \\ere about
eight months old when they were planted for the field trials; lot C, seeds
from the original lot described above, and stored in 1932. Thus it is seen
that pepper plants grO\nJ. in the field in 1945 represented those from very
fresh seeds (lOt A), those from eight-momh-old seeds (lot E), and those
from thirtccn-year-old seeds (lot C), all being from the identical genetic
strain.
Tomato seeds were similar in treatment to the pepper seed c..xcept that
h'l'o original storage condirions were used instead of one. This was to
permit comparison of the field performance of plants from old seeds
which had been stored under fa.ourable and unfavourable conditions, as
"fell as of fresh and old seeds. Seeds of lot C bad been Stored for thirteen
years in open conuiners in the laboratory. Lots A and E were produced
from C in the same manner as described for the corresponding pepper
seeds. Seeds of lot F had been mixed with calcium oxide to remove
one-half of the moisture and then sealed in tin cans and stored at _ 5C.
for thirteen years. Lots D and E were produced from F in the same
manner as lots A and B were produced from C.
In the case of lettUce, as for tomato, seeds from two original storage
conditions served as bases. Both lars had been mixed with calcium oxide
to remove one-half of the moisture, but lot B was stored in the laboratory
while lot D was stored at -S"c. Lots A and C were produced in the
summer of 1944 from plants which had been grown from lors Band D.
respectively.
Seeds of all varieties were planted in soil in flats and placed in the
greenhouse in April, 1945. As germination occurred and the seedlings
grew, they were transplanted to individual pots from which they 'Were
later set out in the field.
The flowers produced by aster and verbena plants were taken as a
measure of the effect of age and storage conditions of the secds from
which they were produced. The weights of fruits produced by pepper
and tomato plants were recorded, and the lettuce heads were weighed
at the time when a majority of them had reached the be.<;t suge for
marketing.
M;TER

Plants from seed-lots A (seyen months old) and B (nine-and-a-half

years old) were set our in the field on 29 A'by 19+5. The planting consisred
of eight ruws with t"..cnry-two plants in each, arranged as follows:

ABBAABB
BAABBAA

,fi

>A, etc.

The flower-heads appearing on !.he phnts were cut from one to three

'42

PLAi~TS

FROM OLD SEEDS

times per week, and a record was kept of the number obtained from each
plant. The first flowers appeared on 18 July. The experiment was terminated on 27 August, at which time most of the plants were dead or
dying. Plants from lot A produced 3,461 flower-heads and those from
lot 3 produced 3,378 flower-heads during the season Cfable XIX). These
totals indicate no significant differences in productivity of plants from
fresh or old seeds. However, if earliness in flowering is considered, the
plants from the old seeds (lot 3) were ahead of those from fresh seeds
(lot A) for the first t'.vo or possibly three weeks. This shows not only in
the totals, but also in the number of flower-heads cut from each row,
though the yield from the rows "iraried in lot A from 327 to 546 flowerheads and in lot B from 268 to 543 flower-heads for the season. There was
considerable variation in the flowering of the plants in different parts of
the field, but lots A and B were equally affected by good or poor growth
conditions.

TABLE XIX
CUMULATIVE WF.f.KLY TOTALS OF ASTER FLOWER-HEADS ClIT DURING
THE GROWIKG-SEASON Ol' I9..lj
Row

,
, ""

,
3

,
,
,

6
7

8
Totals

",

33'

475
3'9

533
366

54-6
38,

37
'9
'5

,,8

176

,,8

323

33'
,,8

'39

3'4

9'

'73

,67

8 ,83

"5

123
'39

25~

Row

""
'"
'4'

"'7

Lot A from seeds 7 months old.

after 1-7 weeks

37 1
346

.....
,,6
378

473

43'
38,

", ""

, ,

,,
"
;

,
6

"7
3'7

1,05 8 1,9,9 2,860 3.344 3,461

47

42
3'4

VERBENA

Lot B from _<eed~ 9'5 )"caJ'S old.

after I~ weeks

8,
3'
66

;8
6;
58
33
53

""

'39

337

'"

199
344

38,
,6,
42 1

432
'95

3#

45 6

335

398

4-41
3"
,6,
543
#9

'4
,6, ,,,

352

~o5

194
25

173

25

365

'36

'"

45 1
5J 5

43 6
''4
,6,
47S

4 24

,68

,8,

1,497 2,398 2,947 3.:190 3.378

Because of the growth habits of these plams and tbe difficulry of

securing exact data on thcir productivity, eighty-eight plants each of lots
A and B were gro\\TI side by side. No observable differences in the ru'o
lots appeared during the growing-season of 1945.
PEPPER

Plants from the three lots of seeds described above wcre set out in the
field on 29 May 1945. For the arrangement of plants in the field a standard
3 X3 form for a Latin square was used, with reshuffling of rows, columns,
and trcanncnts to form ten blocks of nine plants each, consisting of three
plants from each age of seed. The yield, measured in weight of pepper
'43

SEED PRESERVATION AND LONGEVITY

fruits produced in each of the ten blocks, is shown in Table Xx. All
fruits except dIe final harvest were removed from the plants when they
\ycre just beginning to turn red, anti were \\cighcd forthwith. Lots A, n,
and C were produced from fresh, eight-month-old, and thirtecn-ycar-old
seeds, respectively.

TABLE XX
TOTAL PEPPER FRUIT YIELDS IN OUNcr..s FOR SEED-LOTS 11.1\'1) BLOCKS
rOR THE ENTIRE SE.4.S0N
I~t

Age of

seeds

Block

3
m

Fresh
8 mono

6'.'

63

8,

f3 yr.

34

79
93

Totals

5
73

105
53

60

",8

63
59

'78 235 2'4

266

195

52

"

Totals

7
,,8

56 125

59

95

77

39

9'

34

'38 3'4

,"

w
74

82 9

85
87 125

730

308

2,3 l2

297

,,'.

753

It is immediately apparent that the block or position in the field

influenced thc y-ield. A total of only 138 oz. was produced in block 6, for
example, while block 10 produce4 308 oz. of fruit. Differences in yield of
plants from the three seed-lots appeared very small and of doubtful
significance. An analysis of variance test of these data confirmed the above
findings.
The totalities of fruits remaining on the plants were harvested on
22 September 1945, the individual fruits being graded for size and photographed immediately after harvest. It will be seen from Plate I I that the
most large fruits were harvested from lot A, the plants from fresh seeds,
on this date. Weights of these fruits were included in the totals of Table

XX.
A measurement of the heights of the plants made on IS July also failed
to reveal any differences in the lots. A somewhat greater number of fruits
were produced by lot A.
TOMATO

It will be recalled that SL'" lots of tomato seeds were sown for the
measurement of yield as follows: lot A, fresh seeds, and lot B, eightmonth-old seeds-both produced from the original lot C which had been
stored for thirteen years in open containers at room temperature when
the field tests were begun; lot D, fresh seeds, and lot E, eight-month-old
seeds-both produced from the original lot F which had been stored for
thirteen years sealed with calcium oxide at -5C. when the field tests
were begun.
The seedlings were set out in the field on 2 May 1945, except in the
cases of nine plants of lot C which were planted in the field on 12 June.

'44

PLAi\'TS FROM OLD SEEDS

This was necessitated by the failure of the seeds of this lot to give the
expected germination percentage and the consequent need fur a second
planting. These nine plants were thOSe of blocks :2 and 3. Jc may be seen
from their productivity as compared with plants of lot C in the other
blocks (Table X.XI), that the delay in planting had no significant effect.
Seeds of lot C were slow in germinating owing to their germination
capacity being decreased by improper storage. Consequendy the plants
were slower in development than those of the other lots. This may be seen
in Plare r2, which illustra[cs a typical plant of each lot on 22 May when
they ,vere transferred from the greenhouse to the field.

TABLE XXI
TOTAL TOMATO FRUIT YIELDS IN OlJNCf.'i FOR SEED-LOTS AND BLOCKS
FOR TIIE [NlIRE SEASON

Lot

Age of

,""'"

Block

Totals

798

708

543

339

738
47'
66,

59+
,60

3,81 9
3,75
1,845
3,755
3,589
3,9 19
20,632

F=h

5>1

II
C

8 mono
J3 yr.

643

D
E

Fresh

'99
59 2

568
397
"3
650

8mon.
r3 yrt

6"
608

45'
7"

'"

599

30

6,.6

6Bo

478

65 8
599
'7+
635
7'3
795

3,33 1

2,981

4,028

3,857

:q6I

3.674

F
Totals

0,7

5 19

.. Stored in open containers in the bboratoI'y.

t Stored with reduced moisture coment in seJ.!cd containers at - SQC:

Fruits were harvested when ripe and the number of fruits and their
weights were taken for each plant. Pickings were made on twenty-four
dates during the growing-scason. On 25 September, when there was
danger of frost, all of the green fruits remaining were harvested. Their
weights are included in the figures for total )icld.
An examination of the [Otal yield of ripe fruit from each lot (Table
XXI) shows the decidedly inferior production capacity of lot C. Also,
certain parts of the field were better than others for tomato fruit production as indicated by the differences in yield of the blocks. Each block
represents a random reshuffling of rows, columns, and treatments of a
st:tndard 6x 6 form for Latin squares. The six replications of blocks were
equal to the number of treatments-in this case different seed-lots used
for plant production-and also [0 he number of rows. One tomato plant
was placed in each cell of the Larin squares.
An amlysis of variance was made, using the weights of [Ornata fruits
obtained from each plant in the field. The difference in lots was highly
K

145

SEED PRESl.RvATIO~ AND LON(,"EVITY

signifieanL, as was also me difference between Latin squares. A detailed

analysis revealed the source uf the difference in lots. The ~icJd of lot
C versus lot F showed the advantage of a good storage temperature, -5C., combined with drying and sealing, over storage in the
laboratory. Both of these seed-lots were from the same original source,
and both had been stored for thirteen years before they were planted.
Thus it is demonstrated once again that the actual age ofthe seed is o!,much
less impOrllJllce than fhe mvironJ1le1lt ill which it is he/d. Furthermore it was
demonstrated that the vigour of lot C, as measured by the ability to produce high-yielding plants, was decreased. But from the fruits which were
formed, good seeds were obtained (lots A and D). There were no significant differences in the weight of fruits from plants of lots A and D, and
hence fresh and eight-month-old seeds from the same source were
equally good. Thirteen-year-old seeds of lot F, and fresh and eightmonth-old seeds produced from lot F, were all equal in value for use in
field plantings. Thus the total variation in lots is to be accounted for by
the poor yield of lot C.
LEITUCE

Lots Band D represent lettuce seeds stored for thirteen years in the
laboratory and at -5C., respectively. Lots A and C represent fresh seeds
produced from lots Band D, respectively, and stored in the bboratory
for about seven months before planting for field tests. Thus once again in
this experiment we can see the effects of storage conditions of seeds of
the same age upon the plants grown from them. Also, we can compare the
performance of seeds of different ages from the same source.
A standard 4X4 Latin square form served as a basis for the field
planting. Eight replications were made. Two lettuce plants were placed
in each cell of the Latin square. The measure of lettuce plaU( yields
differed from those reported for aster, verbena, pepper, and tomato. In
the four latter cases, flowering and fruiting (i.e. the development of organs
concerned with sexual reproduction) of the plants served for comparison.
But as the commercial value of lettuce depends upon its vegetative growth,
the sizes and weights of its leafy heads were measured. The seeds were of
the variety Iceberg. All of the plants were harvested on the same date,
21 June 1945. The entire plot was screened-to prevent damage by rodents.
A summary of the yield in weight by blocks is presented in Table
XXII. A statistical analysis of these data showed a significant variation in
lots, rows, and columns, and between Latin squares. The cause of the
variation of the lots is strikingly demonstrated. In this case the temperature at which the seeds had been stored for thirteen years was v;ithout
effect on the plants produced from them. The age of the seeds, i.e.
whether they were seven months old or thirteen years old, had a marked
effect and was in favour of the old seeds. It should be pointed out that the
summer of 1945 was a poor growing-season at Yonkers, New York. There

'46

Pl.AIT. II

Complete har"CSt of fruilS R=aining on pepper plants Z2 September, 1<),+5- E:u:h pile of fruits
is 20 inehes in diameter at the ba;;e. Top 10 bottom, lots 1<, R, and C, from plants produced from

fresh, eight-month-old, and thirtccn-ycar-old .>ccds, respccti,-cly.

PL\TE 12

The appearance of tomato plant!< at {he time they were set out in the field on 22 ?Iay, 19.0\5.
Lrfi lu rj~ht .. t]pic:l1J'lants produced from seed-lots ,~, B, C, D, I:, and F. (Lot A, fresh seeds, and
[01 B, cight-month-ol seeds-both produced from the original lot c "'hich had been sw~d for
thineen years in open containers at room tcmptnture when the field {CSIS wen: begun; lot D,
fresh seeds. and lot E, eight-month-old sccd5-both produced from the originaIIOl}' which had been
stored for thirteen }"ean; saloo ....ith calcium oxide at - S-c. when the field tens II'tte begun.)

PLAl\'TS FROM OlD SEEDS

was a great deal of rain and the temperature was low from 29 May to 7
Junc. Behaviour of seedlings in the field might vary \\ ith diAcrent weather
conditions.

TABLE XXlI
TOTAL WEIGlrrs OF LETTUCE PLl\.!':lS, IN OUNCF.5, FOR SEFD-LOTS At'!1)
llL()(XS

Age of

,,""

L"

7 mon .

~T.

13

7 moll.
13 yrt

Totals

Block

"

76

,6
76

88
87

75
93

6,

80

'",

Tl

3"'

355

]06

To"'"

75

,60
574

8]

79
9'

8,
6,

-,.,

65 8

]'"

]"9

316

2.433

, ,

,6
66

7'
86

75
73

,,. '4'" "

75
66

"

7
6,
Tl

6"

Stored with reduced moisture content in setled cont;liners in the bborJ.lory.

t Stored I'~th reduccd moistun: contellt in sealed containers at - S6C

All of the results for aster, verbena, pepper, tomato, and lettuce
reported above were for the behaviour of plants which had been grown
from the variously stored seeds and established in the field. In only one
case, i.e. that of tomato seeds "'hich had been stored in an open container
in the laboratory for thirteen years, did old seeds germinate to form plants
which were inferior in the field. This is not to say that there were no
differences in the germination capacity of fresh and old seeds of the various
plants. The particular lot of tomato seeds just mentioned, for example,
gave only 6 per cent seedling production in soil in the greenhouse. This
is to be compared \\~th 85 to 97 per cent for all of the other lots-including
the ones stored for thirteen years at a low temperature. The germination
of the various lots of seeds was shown to have no direct relation to the
field performance of plants grown from them. However, if the viability
of the seeds is reduced too greatly, delay in germination followed by slow
plant development will decrease their value for the production of seedlings
of good quality, as was seen for tomato seed of lot C. Of course the acmal
number of plants produced from a batch of seeds, as well as the quality
of those plants, is of great importance in commercial practice. If it is
known, though, that good plants will be produced, the density of seeding
can be adjusted to the known germination capacity. The data here presented would certainly justify the use of old seeds within rather widc
age-limits.
As recently as 1951, Schwass (1951) indicated his belief that it is morc
economical in many cases to purchase certified seed of relatively low
germination capacity, and that the resulting crop should be as good as
from seeds giving high gcrmination in the field, provided the so\\~ng
density is increased.
147

CAUSES OF DETERIORATION
D1FFERf.!'H explanations and theories of the cause of deterioration of seeds
have been set forth. Regardless of the cause, ageing seeds result in abnormalities in the seedlings they produce. TIlls is first evident in the delayed
germination of old seeds and is often followed by the production of some
chlorophyll deficiency Or other abnormalities. Increases in the temperature and humidity of storage conditions for omon seeds increased the
number of abnormal seedlings which failed to form sharply bent 'knees'
and possessed blum, undeveloped primary roots (ll. E. Clark, 1948). This
author stated that it was not clear from his results whether the abnormalities were purely anatomical or whether they were due to physiological
deficiencies. It seems safe to say that both anatomical and physiologiCil
response." are involved, but that the mechanism of action has not been
elucidated.

CHE..,,-,uc..u.

Co"'WOSITION

Loss of viability has been thought by many people to result from a

depletion of the food supply of the embryo. However, a cursory examination of just a few non-viable seeds comwces one that there is still plenty
of reserve food left in the seed. A closely related view is that the reserves
in the seed are changed chemically, so that they no longer furnish the
nutritional requirements of the embryo. It has usually been assumed that
any major chemical changes were in the embryo itself. However, Kikuchi
(1954), using the method of transplanting embryos of wheat on to the
endosperm, found that the older the endosperm was, the poorer became
the growth of the transplanted embryo, though the embryo tended to age
more rapidly than the endosperm in individual seeds.
Respiration is one measure of the metabolic activi.ty taking place in
the stored seeds. A low respiratory rate has almost certainly been attained
by seeds that have a long life-span, whether in dry storage or in moist
soil. On the other hand, increased respiration is apt to precede total loss
of viability. The respirawl}' activity of wheat kernels was not-affected by
the length of the storage periods when tests were made after rebtiye1y
short periods of SL":, eighteen, and thirty months (Leach, 1943). Increased
moisture content and low quality of maize seed. increases the respiration
rate, according to Olafson ef uf. (195-1.). Oil-bearing seeds of high moisture
148

CAU$fS OF DETERIORATION

content also have high respiratory rates (Qyam, 1906; Tashiro, IgI3;
Kretovieh & Starodubtseva, 1956; and others). One of the problems in
measuring respiration of seeds is the microbial population which may be
present. For e.xample, the respiration of soya bean seeds containing 185
per cent of moisture, measured after up to three weeks of storage, yielded
curves similar in form to the microbiological population growth-curves
(Milner & Geddes, 194511). Obviously this difficulty is met most frequently in seeds of high moisture content:. Gaseous exchange by seeds of very
low moisture content, on the other hand, is much morc difficult to measure
because of the small quantities of gas involved.
The effect of storage of b'Tain on its carbohydrate and protein content
is of special importance because of the value of these constituents as food.
Wheat grain stored for from three to fifteen years in a cool, dry, wellventilated place in the laboratory was found to be suitable for starch
production, though the recovery of starch decreased somewhat as the
grain aged (MacMasters & Hilbert, 1944). Sugar loss during storage of
different strains of sweet corn seed ,vas about the same from year to year
-indicating that the internal factors effecting the change from sugars to
polysaccharides arc inherited (Doty et af., 1945). There arc small but
definite differences in thc in vitro digestibility of pure starches prepared
from freshly harvested and well-stored seeds of rice, though there is no
marked change in the amounts of the various carbohydrate constituents
(Sreenivasan, 1939). I-Iemicelluloses and proteins decreased, and reducing
sugars and sucrose increased, when expressed as percentage of dry-weight
of blue lupine seeds stored at high relative humidities of 92, 86, and 80
per cent (Ward, 1951). This occurred during the first five monl.hs of
storage and levelled off during the next five months. At lower relative
humidiries of 75,65, and 54 per cent, thc reverse was true, the available
foods decreasing and the storage forms increasing. Depletion of reserve
foods, i.e. hemicelluloses and proteins, was associatcd ",rith loss in viability
of the seeds. Acorns stored under normal atmospheric conditions show a
minor loss of starch and a slight rise of sucrose, with litde loss of viability
(Serenkov & Kuznetsova, 1952). Stored in an atmosphere of carbon
dioxide, the acorns show a loss of starch and a rise of sucrose and monosaccharides, the toral carbohydrates remaining constant.
Crocker & Groves (1915) suggested that the degeneration of seeds in
dry storage may be due to the gradual coagulation of the proteins of the
embryo. Using Buglia's time-temperature fomlllla for coagulation of
proteins, they applied it to the deterioration of wheat seeds at two different
moisture contents and various temperatures, and found that the calculated life-span of the seeds agreed well with the determined longevities
for higher temperatures. Calculated longevities at lower temperatures,
however, wcre much longcr than the life-span of wheat seeds under
ordinary conditior.oS of storage, but they may well conform to the extended
life of wheat grain under the best storage conditions. It remains for

'49

SEED PRESERVATION AI\,'D LONGEVITY

controlled storage of wheat grain to determine whether these calculations

were reasonable.
Three different types of changes occur in the proteins of ground
maize seeds and of whole shelled maize seeds during a storage period of
two years (D. B. Jones ct al., 1942): (1) a decrease in their solubility, (2) a
partial breakdown of the proteins, indicated by a decrease in true protein
content, and (3) a decrease in digestibility. Changes in the ground seeds
were greater than those in the whole shelled seeds, and both were lliOTC
affected at a storage temperature of 76P. than were those stored at 30oP.
Moreover, those in open storage were greater affected than those in sealed
storage. Decrease in protein value and digestibility were more rapid
during early stages of the storage. Essentially the same results were
obtained for soya bean and wheat seeds (D. B. Jones & Gersdorfl:~ 1939,
1941), and for peanut seeds (Moorjani & Bhatia, 1954). Since changes
took place in the flour as well as in the intact grain, the proteins themsel\'es,
whether present in the embryo or the endosperm, arc involved in the
changes.
A study made by Fifield & Robertson (1952) on the milling, baking,
and chemical properties of I\'larquis and Kamed wheat stored for from
nineteen to twenty-seven years, revealed that storage had no consistent
effect on the amount of protein. The best bread was made from the 1921
crop, probably because of its higher protein content.
Rice stored for from forty-six to eighty-four years in granaries in the
northern part of Japan showed decreased amounts of fats in all cases and
of sugars in some; but contents of protein, starch, crude fibre, and ash
were no different from those of fresh seeds, though the power of germination was lost completely (Kondo & Okamura, 1934tl.).
The rather recent reports described above confirm the findings of
Acton (1893), from a comparative analysis of wheat seeds twenty-eight
years old and of new wheat seeds from the same field. In the old sample,
insoluble compounds, such as proteins and starch, had undergone considerable changes to produce substances that were soluble in water. These
changes suggest hydrolysis and enzyme action. The increase in the amount
of dextrin and reducing sugars may have been caused by a slow action of
diastatic enzymes, though no traces of these or of proteolytic enzymes
were found in the old sample. The author believed it probable that such
'fermenL',' were originally present and produced the changes during the
earlier period of -storage, but were later destroyed by m,;dation or hy
micro-organisms. This explanation seems to have been horne out by
later studies which described changes taking place during the early part
of the storage period.
ENZYMES

Because of their importance in metabolism, enzymes hayC been

studied rather extensively in stored seeds, and have been claimed by some

'5

CAUSES OF DETERlOR.o\TION

be an index of the ger"mination capacity and vigour of a seed-lot (see

Chapter XlI). Albo (1908) stated that the loss of germination capa~ty
was direcdy related to enzyme activity. There is no change in enzyme
behaviour in seeds of wheat or their milling products from harvest until
the next spring, wbell dehydrocJenase and peroxidase reached a minimum
and moisture content was at a ma.mnUffi (Kullen, 1941). High storagetemperature promoted, and low storage-tempcramre reduced, the changes
in cnzrme activity-except for catalase, which changed little in any casco
Humidity was also a factor affecting cllZJllle activity.
Amylase activity in rice seeds is greatest at the (milk stage', i.e. during
the maturation of the grain on the plant, and decreases with ad\-ancing
maturity and even more rapidly during the early period of seed storage
(Sreenivasan, 1939; Rao t1 al., 1954). The acn\;ty of amylase of acorns
does not change when the seeds arc stored in moist sand at aC., but
storage in an atmosphere of nitrogen or of carbon dioxide increases the
amylase activity (Evreinova & Erofeev, 1956). Amylase is import:lnt in the
mobilization of starch. in respiration and gromb of the seed.
The catalase content of old seeds, and the possibility of using this as a
measure of viability. has been discussed (Gapter XII). However. it is
unreliable as the results depend upon the seed and the special conditions
under which the measurements were men (Crocker & H:rrrinf,rton, 1918;
Kondo & Okamura, I931, I934)' W. C. Da\"is (1931) pointed out that
catalase and oxidase were frequently used as criteria of metabolism and
viability ofseeds, and that, though the significance ofcat:thse is a question,
the oxidases are definitely associated 'with respiration and other physiological processes which may bring about deterioration. However, he eQuId
find no simple quantitati\"e relation between phenolase acti"ilY and age
or germination capacity; for though young wheat seeds of high germination capabililY showed high phenolase activity, and old seeds of low
germination capability showed low phenolase activity, there was gradual
loss of phenolase activity v,;th age of oat grain-irrespective of viabiliryand this loss continued at the same tatc after comple1e loss of viability.
With cucumber seeds nm, three, and four years old, both viability and
phenolase activity decreased regularly with increasing age, whereas with
barley grain from one to eleven years old, phenolase activity decreased
but little, though gennination dropped from 76 to 12 per ccnt.
Many old seeds "'ere analysed for peroxidase by Brocq-Rousscu &
Gain (1909)' Using seeds of ninety-one genera, they found peroxidase
in I more than So years old; t more than 76 years old; I I more th:l.n 50
ye:l.rs old; 18 more than 25 years old; and 221e5s than 25 years old. They
stated that the maximum known persistence of peroxidase is 20S years.
to

VITAMI!';:;

The vitamin content of old seeds is a subject of some inte1:est. Thiamin

content did not change in wheat grain during short periods of storage
151

SID)

PRESERVATION AND LON(i'EVITY

(Pe:llCC, 19-1-3). Howe..-er, extended storage of from fi..-e months to fifty~ne

years resulted in consistent losses of thiamin with increasing age (Bayfield
& O'Donnell, 19+5). Vitamin B deereased somewhat in rice grain with
storage, but was still present in old seeds (Kondo & Okamura, 1932/33,
1932/330). Increase in the time of storage and in storage temperature
destroyed ascorbic acid in ViglllJ. silletlsis (Hoover, 1955)
AUXTN

The auxin content of maize seeds falls gradually \\~th storage, but
there is still a considerable amount present in twenty-six- to thirty-eightycar-()ld seeds Uuei, 19-1-1). There is no indiCltion that the disappearance
of auxin leads to the death of the seeds.
ALKALOID

The alkaloid content of Slryclmos liIlx-vomica seeds held in a gunny

bag in the humid atmosphere of Debra Dun, India, for sixteen years
remained e$entially the same (Punt:l.mbckar, 19-1-7). It was conduded
that the poor quality of such seeds for the manufacture of the drugs
strychnine and brucine is due mainly to the initial low al.k::tloidal content
of seeds from the same or closely allied species from different localities,
and nOt to any deterioration caused by long storage.
ORG.-\....n C ACIDS

Thl,.'1"e seems to be no doubt that an increase in free fam--acid formation, especially in seeds with high oil contents, accompanies loss of
viabilit), and also loss of commercial value in storage. This has been
demonstrated for corron-seeds and peanuts (F. R. Robertson & Campbell.
1933; Freyer, t9J:l; Hoffpauir ~t ot., I947; Stansbury & Guthrie, I9.i/;
Pons~t a/., 19...8; C. O. Jensen ~t 01.,1951; and others); for grains (Zeleny
& Coleman, 1938; Slusanschi, 1939; Sorgcr-Domenigg ~t al., 1955; and
others); for sunflower (Lishkcvich, 1953); foc Brassica sp. (Tiiufel &
PoWoudek-Fabini, 1954); for fux (Painter & Nesbitt, 1943); for sorghum
and maize (Lindemann, H;J53); for pine (Mirov, 1944); and for Jrlacadamio.
(Chu ~r al., 1953). The fat acidity test has been applied to several hundred
samples of sound and damaged grain, with the result that it has been
possible to establish fat acidity values for grain showing little or no
deterioration (Baker et 01., 1957). It has been shown statistic:illy that
reduction in viability and citric acid content arc parallel in. cereal seeds
(Taufe! & Pohloudek-Fabini, 1955). Deterioration of seeds of i"ledicago
salroa, Trifolium prareme, and LoWS (ornimlatus is due to an accumulation
of lactic acid, according to Wyttenbach (1955). This lactic acid, probably
produced as a result of intensified respirarion, exercises a tm.ic effect on
the embryonic tissues as soon as its concentration is increased. The dead
grllins contained at least fifty times as much lactic acid as fresh gra.ins
having a high germination capacity.
15 2

CAUSES OF DETERIORATION

INHIBITING SUBSTANCF.s

Schwemmle (1940) offered some e."\:perimenrnl evidence of the

accumuhtion of inhibiting or toxic materials in seeds as they age. The
seeds of Oroothera berta-iana germinate more slowly and in lower percentlges as they become older, up to eleven years. By running tests with

fresh sCt.'t!s which were treated with "'Iter extract or watcr-cxtraeted mash
of old seeds, he was able to show that the old seeds contained germinationinhibiting substances. The mash of the old seeds was especially effective
in inhibiting the germination and later seedling growth of new seeds.
Omothuo adora/a seeds also lost gennination capacity with age, but the
development of the seedlings was much less inhibited by extracts of its
old seeds than was the case wir.h O. berteriona. Stubbe (1935. 1935a) has
suggested that the metabolic changes of seeds in dry storage lead to tbe
accumulation of inhibiting or lOne subst::mces that transform the nuclear
materials and lead to mutations.
Artificial mutations of seedlings of Nit:otia~ tablKUm produced from
normal seeds were induced b)' treating them with cold water extracts of
ground, prematurely aged seeds. Oil and oily emulsions of the aged seeds
had considerably less effect than Lhe aqueous extract.. Radiosensitivity of
wheat seeds increases with increasing age, according to Nilan & Guntbardt (1956). The greater sensitivity of the chromosomes in aged seeds
may be due to chemical compounds produced b)' the decomposition of
food reser....es.

l\'IurATlOxs
There is a considerable amount of evidence that disarrangement
of the nuclear mechanism is responsible for deterioration, or results
directly from physiological changes which initiate the deterioration process. Shkvarniko.... (1937) found an increased mutation tate in plants from
six- to ren-year-old seeds of summer wheats as compared with fresh seeds.
lViut:ltions from old seeds showed chlorophyll deficiencies, thick heads,
square heads, speltoid forms, dwarfs, and sterile plants. However,
Shl.,,-'anlikov also found great variation in the mutation ratc from secus of
the same age. It has been shown that seeds of Datura which were aged by
storage in the laboratory for periods of up to ten years, have increased
mutation rates for pollen-abortion mutations, and that these mutations
arc inherited (Cartledge & Blakeslee, 1934).
This wurk followed the discovery by Navashin (1933) that in root tips
uf plants 6'Town from aged seeds of Crepis, chromosomal abnormalities
were produced in large numbers. Shortly later Peto (1933) also found
large numbers of chromosomal mutations in root tips of maize from aged
seeds. The work on Datura has been e}.1:cnded by showing that lligh tates
of visible mutations are al<>o induced from old seeds (Blakeslee & Avery,
1934; Avery & Blaskcslee, 1936), and that the ageing of pollen grains for
periods uf up to thirteen days is even more effective for increasing the

'53

SIn)

PRLSERv,\TION .\;'ffi LONGE\'llY

pollen-abortion mutations than is ageing of seeds (Curledge et al., 1935).

Stubbe (1935) reported that ageing of seeds of AntirrhimmJ grearly
increases the incidence of visible mutations.
These results raise the question of the causes of the increased mutation
rates follo" ing ageing. That the increases arc not due to age alone is
indicated by the results of study of Datura plants grown from seeds that
had been aged for twenty-two years in the soil (Cartledge & "Blakeslee,
1935). These plants showed rates of pollen-abonion mutation that werc
onlr slightly higher than the rates found for their controls. Investigation
of the effects on the mutation rate of varying the environmental factors to
whieh the seeds might be subjected was thus suggested. Peru (1933) has
shown for barley, and Navashin & Shkvarmkov (1933) and Shl'varnikov
& Navashin (1935) fOT CrepiJ, that heating seeds is more effccri\'e than
ageing for the production of chromosorn.al mutations in the root tips.
.o\lthough many other accounts of experiments on hearing seeds may be
found in the literature, they secm to be connected with the control of
fungal or bacterial parasitcs, or with thc effects of the treatment upon thc
gcrminability of the sccds---OT. in a few cases, with the abnormal growth
of the seedlings dC"l'"e1oped from heated seeds.
When the moisture COntent is very low, Sl.WS of m:lnr sorts will endure
surprisingl}' high temperaru:res. Harrington & Crocker (1918) germinated
seeds of Johnson grass that had been heatcd for SL">: hours at 100"C. \\;th
moisture contents of from zo per cent to 01 per cent, but the seedlings
produced were less vigorous than their controls. Waggoner (1917)
reported that q. per cent of somc seeds of the Icicle radish were viable
after heating for ~. minutes at 123"C. V\;th 0'4 per cent moisture content, but that all died which were heated at lOO"C. with 4 per cent
moisture, or at 6S"C. with 40 per eent moisrure. By pre-heatin~ for one
day at from 65" to 7S"C., and for one day at 9O"C., Dixon (1901) was able
to germinate seeds of a number of plant species after heatiD~ them at
temperarures ranging from 100" to Il4"C. AtarulSOff & Johnson (1920)
heated well-dried seeds of wheat, barlcy, oats, and rye at lOO"C. for
fifteen and thirty hours, md of wheat and barley at from 100" to IlO"C.
for fony-five hours, and got good gcrmination--excepr in the case of the
rye seed, which germinated poorly. brlier literature on heating seeds is
reviewed in the papers of Waggoner (1917) and of Atanasoff & Johnson
(19 20).
Waggoner, and other investigators, havc noted that heating of seeds
may cause delay in theu- germination. Crocker & Groves (1915) found
that wheat seeds delayed the beginning of their germination from four to
fourteen days, depending upon the length of heating. Jozefowicz (1930)
found that increasingly severe heat-treatments of tomara seeds caused
progressivc delay in their germination, whieh occurred from ten to
eighteen days af[cr planting. Scedlin:,"S grown from seeds tha[ have been
subjected to severe heat-treatments may show abnormalities of gro""th

154

CAUSF.5 OF DETERIORATION

and form. Jozefowicz (1930) obtained from 50 to 100 per cent of abnormal
tomato seedlings from seeds heated at 90" to 1oo"C. for one hour, unless
the seeds were carefully pre-tlried. Gain (1924) grew sunflower plants
from dcsicr:uted embryos which had endured a series of successively more
severe heat-treatments, ending with tcmperature..<; raised from 125 to
155C. during about thirty minutes. The plants which were grown from
embryos that had been so treated were very abnormal in form and growth,
and although some of them produced flowers, none was able to form seeds
(Goin, '927).
A dctniled experiment in which heat and moisture were invcstigated
as factors in the increased mutation ratc from Datura seeds, has been
described by C1rtlcdge et al. (1936). The seeds used ,vere produced by
self-pollinating Datura. plants of a standard line which had been inbred
since 1916, and had been passed through a haploid phase containing
maternal chromosomes only_ These seeds were subjected to treatments
with various environmental factors, particularly 'with heat and controlled
moisture content. Mter adjustment of the moisture contcnt of the seeds
to 2 to 15 per cent, they were placed in sealed vials and treated at temperatures of from 45(> to 80C. for from two hours to five days. The
largest numbers wcrc tre..1.ted at 5 pcr ccnt moisrure and 75 to 80C. for
from two to forty-eight hours. The most severe treatmcnts killed the
seeds, e..<;peciallr whcn these were high in moisture content, but modcrate
treatments increascd the seedling yield O\'cr that of the controls. Good
germination was obtained from treatments of seeds wi.th 5 per cent
moisture for twenty-four hours at 80C., ana for thirty-six hours at 75C.
The interval of time bctween planting and appearance of the seedlings
was lengthened by increases in the temperature of tre-atment, by increases
in moisture content of the seed, and especiall)' by increases in duration of
the treatment. Seeds with moderate and severe heat-treatments produced
high percentages of plants with abnormal growth, as reflected in their
types of branching.
The plants wcrc tested for pollen-abortion mutations by microscopic
examinations of pollen samples. Mutations of the pollcn-abortion gene
typc were found in fifty-six plants, while pollen-abortion of the type
caused by chromosomal mutations occurred in thirty-seven plants.
Although a total of nincty-three pollen-abortion mutations WCfC thus
found i.n the 8,741 plants tested, none of thesc was found in tbe 920
control plants included. Most of the mutations involved a sector comprising half, or less than half, of the plant. Within the exposure timcs
used, there was no significant increase of mutations at temperatures lower
than 70C. The highest mutation rates found wefe about 5 per cent,
obtained from seeds with 5 per cent moisrure content that had been
heated at 75" and SaC. Higher rales had been obtained from aged seeds
(Cartledge & Blakeslee, 1934). In general, the mutation rate increased
with increased temperature, with increased moisrure content, and with
155

SEED PRESERVATION AND LONGEV!TY

increased duration of ~eatmcnt----<:onditionswhich dcla~'ed the germination of treated seeds.

The primary purpose of the experiments by Cartledge ~t al. (1936)
was to test the effects of treatments which had been selected as possible
factors in the increased mutation rates obtained from aged seed.
The effect on Ute mutation rate of en\-ironmental factors such as heat
and moisture, is of direct interest for the problem of the cause and nature
of mutation. Although, not SO many years ago, it was'generallr supposed
that mutation was both rare in n:tture and beyond the reach of experimental stimulation, it is now well kno~"TI that a variety of experimental
procedures may be successfully employed to induce mut~ttions. Those
recognized at first were strong radiation treatments; but it has become
increasingly apparent that treatments more nearly related to the normal
environment of the organism arc capable of producing similar, though
less extreme, mutational effects. Heat is a factor in the normal em~ron
mem, and one to which cclls respond by quantitative or qualitative
chan"ores in their physiological processes. Kavasbin ('933) has suggested
that age and heat treatments uperate to produce mutations through
alteration of the rate or character of the normal processes within the living
cells. If this should be the case, then the mutations induced by a1!e, hot,
and the like, may be related in cause to those mutations considered natural
or spontaneous. Increased mutation rates from ageing seeds of Datura
(Canledgc & Blakeslee, 193+) may be explained as an accumulation of
mutations occurring at a more or less constant rate. Stubbe (1935, 19350)
presents evidence for an accelerated rate of mutation ~;th age in seeds of
Al1tirrltinum. Heat, on the other b:l.I1d, certainly increases the rate of
mutation; but the effects of age and of heat may be reducible to a timetemperature relationship ~hich \\ ill include both treatments.
It is of some interest to observe that tbe experimental detcrminations
of normal mutation rates arc usually made under carefully controlled
conditions ~;th fresh material. The increases of mutation associated 'tT-ith
ageing, high moisture content, and heating of the seeds, suggest that
condition... bcyond the-limits of carefully controlled experiments may be
of importance in determining mutatiOIl rates in nature, and that these
ratcs m.ay be higher than the 'normal' rates found by experimental tCSts.
It is obvious from the results of GnIedge et 01. (1936) that the
maximum mutation rate precedes the death of the seeds , as the treatments
giving the highcst pollen-abortion rates still permitted excellent germination. This was also noted by Navashin el 01. (1940), who found that the
life of tbe seed of Crepis was proportional to the time of storage, but the
mutation fate was proportional to the time only in the early storage period,
i.e. the flrst 1,000 days. After 1,000 days a ,~-fold increase in the storage
period resulted in nearl~' nine tim(S more mutations.
More recent work has served to emphasize the increased mutation
rate with ageing of different seeds. Mutants characterized by abnormal
t5 6

CAUSf.,> OF DETERIORATION

height and increased number ofleaves appeared as a result of ageing seeds

of Nicol ill/Ill fubuClII/I (Gisquet et aI., 1951). The fOOL" oflcttuce and onion
seedlings developed from old secds show much chromosome breakage,
resulting in bridges and fragments at ana.phase (Lewis, 1953). Harrison
& McLeish (1954), working also with lettuce and onion roots, found a
general trend to a sharp decrease in chromosomc abnormalities at high
germination values, and a corresponding rise in mitotic activity. Results
with onion, however, revealed a low incidence of chromosome breakage
at all levels of germination, showing dlJlt loss of viability is not necessarily
determincd by, nor does it precede, increase in the frequency of abnormal
cells. These authors conclude (Harrison & McLeish, 195--1-) that the production of cytological abnOImalities must be a consequence rather than
thc initial cause of seed deterioration.
Cytological and genetical changes occurring in ageing seeds of common
and dUnlm wheat, barley, rye, and peas, that had been stored from one to
thirty-two years. have been studied by GUlldlardt et al. (1953). Their
analysis included a record of (1) chromosomal bridges and fragments in
mitotic :maphase of root-tip and shoot-tip cells of seeds of various ages,
(z) chromosomal interchanges in the pollen-mother cells of plants grown
from these seeds, and (3) seedling mutations in the progeny of these seeds.
Examples of chromosomal aberrations are shown in Plate 13 (Gunthardt
a ai., 1953). Both chromosomal aberrations and genetic mutations arc
described, and the types of the former appear to be identical to those
arising from ionizing radiations. The authols also :;how that the dosage of
natural radiation, including cosmic radiation, is not sufficient to account
for the changes taking place in the ageing seeds.
Gustafsson (1937), in his description of death as a nuclear process,
s:tys that vital processes are going on in the nuclei of the embryo in the
seed preparatory to the reproduction of chromosomes and nuclei. If these
processes have advanced to a certain stage, but e-x1:ernal conditions
prevent their fulfilment in the actual division, the nuclei degenerate and
die. Inereascd moisture content hastens this stage. and thus shortens the
life of the seed.
Amato (195+) has critically reviewed much of the literature on physiological and genetic aspects of seed ageing in connection with the problem
of spontaneous mutability in plams, and has given an extensive bibliography on the subject.
A<; storage conditions can overcome the effect of ageing, as measured
by rears of life, a ncw definition for ageing in seeds is needed. Longevity
is more closely related to the morphological and physiological state of the
seed, especially of the embryo, than to dIe length of the storage period.
'Ageing' takes place in a few hours under some conditions, while under
optimum conditions ageing might require centuries. More basic research
is needed on this problem, using such new 'tools' as chromatography and
radioactivc tracers, to detennine the basic causes of loss of viability.

157

XVI
PRACTICAL CONSIDER->,TIONS
TRANSPORTATION

1'HF.RE is a good chance that seeds, especially those which are \"ery
sensitive to high humidity and temperatures, will lose their viability in
transit. Large consignments of field seeds could be killed by overbeating
in railway trucks or the holds of ships. The fIrst thought is that such
deterioration could be prevented by drying the seeds and dispatching
them in scaled containers. This ,",auld probably be effective for most
seeds, but there still remain some u-hich do flot tolerate desiCC:l.tion ;lod
must be kept moist on the way_ This is a difficulty in transporting palm
seeds. However. a satisfactory method of packing palm seed for distant
transportation has been developed at the Puerto Rico Experiment Swion

(Anon., I94ob). Experience at the station suggested that the failure of

palm seeds to germinate may have been the result of improper packing _
of the seeds at their SOUTce. As the seeds must bl.: kept moist, they arc
often made tOO wet and hence germinate, the resultant seedlings dying in
transit. E.."\:ccUent results can be obtained by packing the seeds in a
medium of granulated peat-moss or the dusty, granular, corky lll:lterial
from coil', in either case having a moisturc content of approximately 30
per CCnt. The mi.'\ture of seeds and packing material should be hermetically sealed in ordinary tin OIlS. If the proper moisture content has
been obtained for the pacl:i.ng material, and the tin container is properl:"
sealed, the palm seeds will not lose sufficient moisture to lmlcr their
viability, while on the other hand they cannot absorb sufficient moisture
to germinate during tranSit.
This problem has been overcome in a different way by Kurokami t:f Ill.
(19+7). It has been the practice to pack seeds of chesmut, which are killed
by drying, in moistened Sawdust for storage or carriage. This is a good
plan for storage at low temperatures, for germination does not take place
or is, at least, delayed. However, during storage at high temperatures.
such as would be encountered in rransit, there is danger of germination
and subsequent loss. These authors inhibited such germination by using
2 to 4 gm. of the phytohormone, naphthalene potassium acetate, or ilS
mcthyl ester, per packing box. The boxes were approximately I ft. 8 in.
long, 1 ft. wide, and I fro deep. Sawdust layers at the top and honom ",ere
about +in. in thid:ness. The phytohormones were prepared in aqueous

15 8

'
,r.~

..j

.
"*,
,,---.)

Photo. byrollrlr5Y W/l,hingl"n Agri"dt"'Il{ Ex~rjmtnl Sia/ion, Pllllnwn,

Wilshingtofl, U.S.A.
PUT~ 13

Examples of chromosomal aberrations ocrurring in plants from

aged ....heat seeds. A, Kub:mh 1(119: llridge in mitotic anaphase,
X 900. B, Kuh:lllka 1919: Ring-of-four in mitotic mC12phasc, X 900.
c. Kuhanka 19'9: Bridge lod:cd into a diccntric chromosome, x Jo8o.
P, Haut 192j": Two parallel bridges in mitoue anaphase, X lo.~O.
E, Kub:mb. IllIg: Two interlocked bridges in mitotic :lrutphasc,
X 1080. P, BaaTt 1925: Two LT<)S$OO bridges, one pair of acentric rod
fra.nncnts, and fWD acentric rings, in mitotic anaphase, X TOSO.
G, &:;;! 1925: Nine bridges and sc\'Cral fr:Igments (some pa~d) in
mitotic anaphase, x 1080. (From Guntharrlt, Smith, Haferkamp &
Nilan, 1953.)

PRACTICAL CONSIDERATIONS

solution, which was sprinkled over the sawdust "\\~th a watering can and
then mi);cd into the sawdust by hand. Depending on the market price of
plant hormones, this method can be put to practical use in the storage of
chestnut seeds over cnended periods of time, or in the transport of these
seeds over long distances through sub-tropical regions (Kutokami et aI.,

1947)
In J941, a ton of selected rubber seed was flown to Brazil in three

large army bombers (Anon., I94Ih). The seed was collected from highyielding clone.,> gmwn in the Philippines, and was shipped to the Canal
Zone by boat. Delay in reaching the Canal Zone made it imperative to
deliver the seeds quickly to prevent dcrcrioratiOll_ For small quantities of
valuable seeds, the aeroplane offers a method of maintaining viability in
transportation, but it is not yet practical for large seed-lots.
It is the opinion of Degen & Puttemans (1931) that seeds do not lose
their gcnnination power as a result of shipment across the ocean or the
Equator, but rather in storage in tropical countries. Farmers and horticulturists living in tropical countries and buying their seeds in Europe,
!>hould secure them as short a time as possible before they are to be used.
These twO authors., Degen in Brw.i1 and Puttemans in Hungary, exchanged samples of seeds from di.IIerent phint families and variously
sensitive to climatic changes, before arriving at the above conclusions.
aearly some seeds are killed by transit conditions while others are not.
More knowledge is needed of the tolerances of various seeds to transport
conditions.
STORAGE FAOLll'JfS

Governments of various countries, in additiun to individuals in

responsible agricultural positions, are becoming increasingly a\\ are of the
possibilities of retaining both fund and seeding value of seeds by proper
storage.
In India., for example, rice grain is more subject to deterioration than
that of any other cereal crop, for it is grown and stored under humid and
often primitive conditions (Panikkar, 1947). About 317.00 tons, 1'5 per
cent of the annual crop. is lost in storage-:J. loss important to a people
living near the subsistence level. India also must import a large quantity
of vegetable seeds each year, because of the short life-span of such seeds
in the Indian climate (Dutt & Thakurta, 1953-55). More thought and
experimental data are being applied to a practical solution of this problem,
so that surplus stocks of vegetable seeds will not have to be wasted for lack
of proper storage facilities.
A survey has been made of the problem associated with the storage of
grain and the methods used in Centrnl and East Africa, with a view to
recommendations of mcans to prevent storage losses (Oxley, 1950).
Although the best condition for maintaining viability is obtained
through control of both temperature and humidity, it Ita!> been found

'59

SEED PRESERVATION AI\"'D LaNGEVIn'

that seed can be preserved for a reasonable period at atmospheric temperature in Puerto Rico, pro\'ided the rclau\-c humidity is reduced w
20 per cent or less (Hopbins (I a/., 19-17). A practical and cheap dehydrating agenr was discovered in ordinary clay subsoil which had been
oven-dried to remQ\-e the hygroscopic moisture. As a result of these
findings, practical storage methods can be recommended.
For commercial purposes in cooler, less humid climates such as those
of New South Wales (Myers, 1943) or Denmark (Dorph-Pctersen, 1928),
110 special storage is necessary if the seeds are to be kept for a short
period, i.e. one or two years.
1\. cold-stomge room is an excellent place to store large quantities of
seeds, according to Akamine (J943), though care must be taken not to
aUow the drip from melting ice to come in contact with the seeds. The
seeds should be placed in the driest pan of the cold-storage room. Other
work has shown the danger of a cold-storage room y;;th a humid atnlOsphere, unless the temperature is below free-ang-point.
Weibull's data (Wei bull, 1955) also show the possibility of using a
commen:i.al cold-storage facility, but he recommends an operating tetnperature of -20"C. for long-term stonge of certain vegetable and flower
seeds. He points out that, v.;th cold storage, the seed trade could be made
less risky and the supply of seeds more regular; prices could be kept at a
more even leveL, and thc plant breeder could concentT:Itc more efficiently
upon his stock-seed production and accumulate bigger stock-seed
reserves.
In the United States as of 1954 (Anon., J954), storage losses of cereals
(wheat, oats, b::tIle~', rye, and rice grain) amounted to the harvest from
3,676,000 acres, "alued at &},666,ooo, and representing a ~'5 per cent
loss. This was exclusive of losses due to insects, which accounted for a
loss of the production from an additional 1,428,000 acres, representing a
loss of 2'4 per cent of thc crop. Recommendations for practical dt};ng
procedures and storage facilities have been made for specific crop seeds
in the various states; i.e. in Texas for rice (Hildrclh & Sorenson, 1957), in
Illinois for soya beans (D. G. c"mer & I Iolman, 1952), in Kansas for wheat
U L. Schmidt, 1955), and in South Carolina for cotton (Brand & Sherman, 19t3). Nebraska has prO\;ded a source of pure seed both of newly
developed varieties and of the more important established varieties (Sahs,
1957). The federal soil conservation service is providing nursery storage
cellars at Pullman, Washington, with air conditioning, to help preservc
tree seeds o,,"er long periods of time (Anon., 1940). A national seed stor..ge
facility where seed stocks will be preserved for future use has been constructed on the campus of Colorado State University at Fort Collins,
Colorado (Anon., J958; Binkley, 1958). It will scrve as a federal germplasm bank on which plant breeders can draw for work ~;th tholls:mds of
different plants representing the world's most valuable food, feed, pasture,
fibre, and tree crops. It is hoped that this will prevent the loss of many

,60

PItACflCAL CO:-lSIDERATJONS

r.ew desirable forms as well as old varieties during years of active plant
breeding. Sccd stock from all ovcr the world will be stored for future
refercnce, and a seed-testing laboratory will be part of this new building.
Descriptions of seed storage rooms, prnctices, and equipment, as
employed in various localities, are to be found in the following articles:
Anon., 1931; Long & Cropsey, 1941j Cartter, 19+2; Eggleston, 1949;
Singley ct aI., 1954; Swainc, 1954; and Pingale & DaIu, 1955
Recommendations such as have been outlined in tIlls chaptcr could
not have been made without numerous experiments to determine the
specific requirements for safe storage of various kinds of sccds. They
represent a great advance in seed technology. At least two experiments
arc now in progress to obtain detailed information on seeds over longer
storage periods. One of these has heen described by Went (1948) to last
a possible 360 years. The other involves lettuce, onion, and tomato seeds,
in sufficient quantity for 150 years of testing, which have bccn stored at
the Boyce Thompson Institute for Plant Research, Inc., Yonkers, New
York (unpublished). As, in this latter instance, dry seeds arc stored in
sealed containers at a temperature of -17"gC., it may wcll be that some
of the seeds will still be viable at the end of 150 years.

GLOSSARY
Accessory fruiL One in which some moo p:u1 of the phnt not bdonging to
the pistil (gynaecium) is associ:l.\ed with the: nurun:d O\1Uy.
Achenc. A small, dry, ane-sccded indchiscem fruit.
Actinomycytes. A group of filamentous organisms rehu:d [0 baeteri:l..
Aerobic. IJ\ing or actiH: only in the presence of ~-gcn_
Aggregate fruiL A fruit formed from SC\"era! o\"2rics produced by one (lov.'Cr.
Alkaloid. AD organic substmcc occurring Il2tul'llll)' in phnts, e.g. morphine:.
Am.)'Jase. An enzyme accelerating decomposition of srarch.
Analysis of variance. A statistical treatment of experimental results.
AnaphllSC. A mgt in cd.l di\ ision during divergence of dau~hIcr chromo.<;omes..
Anatomy. The scic:nce which IrC2ts of suucnm: or, more ""idely, of form. Plant
anarom~ deals with the cdls which make up pb-nlS.
Angiosperms. Plants with seeds borne wilhin the OV':lry.
AlllluaJ. Of only one year's duration, or less.
Anther. The pollcn-bearing ponion of a mmcn.
Aquatic. Living: in w-.ner.
Ascorbic acid. Pure Vitamin C.
Aurin. Growt.h-regularing hormone.
Biochemistry. The chemistry of hing organisms :md their components and
prouucts.
Calyx. The outer ,... hor! of floral 'le:I\"tS'.
Carbohydrates. St:lrChcs md sugars.
G.rpeL A simple pistil or one member of:l compound pistil.
Catalase.An enzyrlle IoI-hich decomposeshydrogcn peroxide into W;lter amI Olt:ygen.
Chromosome. A dt'eply-staining body, normally of a constant number for one
species, found in the nucleus during edt division.
Oone. A group of pl.an.rs der1,-ed from a single plant by vegetative prop:!gation
or artificial sepantian..
Cot)'lwon. The foliar ponjoo or first leavcs (one, rn-o, or occasiona1ly more) of
the embryo as found in the seed.
Critjcal moisture content. Seed moisture content above which dcterior:nion is
npid.
Crude fibre. Material obtained as residue in the ebemicol aJUl)"Sis of plant
substaIKCS.
Cuticle. The thin, usually "Ilo'<lJty oo\ering of Ihe outside of epidenn:ll cells of most
land plants.
Cytological. Perl.aining [0 the celL
DchiscenL Opening sponuneously along I,;crt:l.in lines or .in:l definite direction.
Dehydrogenase. An enzyme accder.ning the rcII1O\-aI of hydrogen from lis5ucs.
Desiccation. Drying out.
Dextrin. A soluble substance dcri\'ed from starch.
Diaphorase. A dehydrogenase <:ont:tining ribofla\w.
Diastase. An enzyme which acts principally in converting swch into sugar.
Dicotyledons. Plants M\ing seeds \l,ilh two cotyledons or secd-It:l.\"cs,:md c0mprising: a group induding most of OUT f:uuili:tr broad-leafed trees and \'egetablts, elc.
162

GLOSSARY

Drupe. .A fleshy fruit with lhe inner portion of the pcricarp h:lrd or ~ony.
Electrolytc. A nmduetor of electricity.
Electromoti\oc [orce. One cau.',ing ekctrica.l acrion or enects.
Embryo. The rudimem::rry phntlet within the st:ctl.
Endosperm. The nutritive tissue of certain seeds.
Enzyme. A chemical acting on one or more specific substrates; a terment.
Epidermis. The outside layer of cells of the aerial parts of higher plants.
Ferment. An en7;}'IJle.
Fertilization. Fusion of m:tIe and female nuclei to form the begirmillgs of a new
individual.
Fluorescence. The property of emitting radiation.
Fruit. The seed-bearing portion of the plant, formed from the ripened ovary
together with other pl::mt parts which may be associated with it.
Fungi. A grcat group of lower plants without chlorophyll, and typically saprophytic or parasitic. They cause many plant diseases.
Fungicide. A substance which destroys fungi.
Galvanometer. An instrument for measuring a small electric current.
Gene. A unit hcrcdit:ll} mcror in the chromosome:
Genetics. The bnuch of biology dealing 'with heredity and variation.
Grain. The fruit of cereals.
Gynmosperms. Plants bearing naked seeds without an ovary.
Haploid. Organism (or state of) having the number of ehromooomcs characteristic of one mature germ-cc11.
Hcnlicellulose. One of SCn':ral pol)s:l.ccharides occurring a.~ constituents of cen
walls, cotyledons, end05pcrms, and woody tissues.
Hermetically sealed. Air-tight.
Hespcridium. A bcrry covered \yirh a tough rind, as in orange.
Homozygous. Having identical genes for a gi'-en chancter.
H)'brid. A cross-breeJ of two species (normally).
Hydrolysis. Reaction between a compound and watcr. .
Hygroscopic. Sensiti,'e to or retaining moisture.
Hypocotyl. Parr ofthe embryo between the cotyledon and rhe radical in the secd.
Imbibed secds. Seeds which h:l\'e absorbeclliquid.
Indeblscent. Not opening or splitting at maturity.
Insecticide. A substance which kills insects.
in pitro. In :l test-tube; outside of the orglmism.
Jacobsen germinator. A special device for seed testing in which water i.~ supplied
to a blotting-paper substrate by mean.~ of wicls.
Latin square. An experimental design comaining the same number of replications
and treatments.
Lignification. Thickening of plant cen-walls by deposition of wood.
Longevity. Life-span.
Metabolism. The chemical changes, constructive and destructivc, occurring in
living organisms.
Microflora. Microscopic pla.nts, such as fungi.
Miero-organisms.l\'licroscopic org:m:isms.
Mitosis. Nuclt::lr division.
Monocotyledons. Plants having seeds with one cotyledon or seed-leaf, and
fanning a group (including grasses, palms, etc.) that with the dicotyledons (q.v.) make
up tile Angiosperms (q.v.).
Monosaccharide. A simple sugar, not decomposed by hydrolysis.
Morphulogy. Thc science of (mostly gross extemal) form and structurc of living
organisms.

SEED PRESERVATION Ai\,TJ) LONGEVln"

Multiple fruit. One resulting: from !.he union or compact aggregation of o~aril"S

or lllore tb:m one flower.

Mutation. Varia,ion producing :I. dcfiuiLt, inhcrcm amI heritable change in an

organism.
Necrotic tissues. Dead tissues.
Nicotinic acid. The pellagra-preventive facmr of Vitamin B comJ:?lex.
Nucleus. 111e complex and usually spheroidal mass e<>smtial to the life Qf most
cells.
Nut. A hard, indehiscent, onc-seeded fruir.
Organic acid. An acid containing at least the elements carbon, oxygen, and
hydrogen, and nonnally produced by or derived from tissues ofliving organisms.
Ovary. The part of we pistil (gynaecium) which contain.__ the ovules.
Ovule. The body which becomes the sccrl after fertilization.
Oxidase. An enzyme- which promotes o:ciilition.
Palisade cells. An arrangement of elongated cells 10 form a tissue, usually constituting the uppermost green layer in the leaf.
Pathogen. An organism causing a disease.
Pepo. A fleshy frui[ with a hard rind, e.g. a squash or melon.
Perennial. Penasting for a number of years.
Pcricarp. The illatured ovary-wall.
Pcrmeahiliry. State of being penetrable.
Peroxidase. An eilZyme which causes activation of peroxide oxyg~n.
Phenolase. An enzyme that oxidizes phenols.
Ph~'si()logy. The science of function and activity of organisms.
PhytohorInones. Internal secretions of plants; plant hormones.
Pistil (Gynaeeium). The seed-bearing org:m or orgam of the flower consisting
of ovary, J;tyle (sometimes absent), and stigma.
Plasmolysis. \Vithdrawal ofwater from plant' cells, causing a comrJction of cell
contents.
Plumule. The part of he embryo which forms the shoot of the plant.
Pollen. The 'male' grains contained in the amher.
Pollination. The process of depositing pollen upon the stigma.
Polyethylene. A plastic film.
Polyploid. A plant with more than two.sets of chromosomes.
Polysaccharide. A carbobyclnte decomposable by hydrolysis into two or more
simple sugars or monosaccharides.
Pome. An indehiscent, two- or more-celled fleshy fru,ir, as in apple.
Protein. Albuminous substance.
Proteolytic enzJmcs. Those which bring about decomposition of protein.
Radicle. The portion of the embryo which forms the root of the new plam.
Radiocarbon dating. A method of age-determination based on the amount of
radioactive carbon found in the sample, as compared with the amount of such carbon
known to be present in young tissues oflhe same type.
Radiograph. i\.ll instrumem fill' measuring and recorrling mdiation.
Receptacle. The more or less expanded portion of a plant bearing the flower-parts.
Reducing sugar. A mono- or di-saceharide that reduces (copper or silver salts)
in alkaline solution.
Relative hUIIlidity. Ratio of the quantity of v:lpour actually presem in the air
to the greatest amount it can contain at a given temperature.
Respiration. C'r.lSCOUS interchange between an organism and its surrounding
llli:<liumj normally the taking in of oxygen and the giving out of carbon dio~dc.
Ribothwin. Vitamin B 2
Samara. An indehiscent, winged fruit, as in ash.

,64

"=

GLOSSARY

Saprophyte. A plant which livcs on dcad or decaying org-.lIlic maLLCf.

Scutell=. Cotyledon peculiar to the grasses, beillg a shicld-shapeJ Of',;an
through which thl: embryo absorbs food from the endosperm.
Seed. The ripened oYule, consisting of the embryo and coats, with often some
additional stored food.
Silica gel. A form of colloidal silica possessing many fine pores, and hence vcr}"
abSQIbent.
Simple fruit. One resultillg from the ripening of a single ovary.
Spermatophyte. Plant producing seeds; a ~-.:ed-plam.
Stratification. Low-tt:mpcraQlre pretreatment of seeds-originally gi\-en 'with
layers of seeds alternating '\\ith layers of sand or soil.
Style. ~111e usually elongated pardon of the pistil (gynaecium) connecting the
stigma and ovary.
Substrate. A substance upon which something grows or an enzyme aelS.
Sucrose. Cane sugar.
Taxonomy. The science of pbm and animal classification.
Thiamin. Vitamin B l .
Topographic. Dcscripti\'e of a plaec.
Vapour pressurc. The pressure of a confined body of vapour.
Viability. The condition of being viable in the sense of being l:apable of growth
and survival,
Vinyl film. A special rype of plastic.
Vitality. Speed:rnd energy of growth.
Vitamin. Acce'>SOry food factor.

165

BIBLIOGRAPHY'
AIlBOTf, E. V. (1950) Some observations on sugarcane flowering and seed productioll
in LouL<;iana and:m experiment on production and storage of true sugar=e seed.
Sug. Bull. N. Orl~Q"S, 28(:u), 32()-32. (Abstr. in Bioi. Abstr., 25, No. 25006,
1951.)
.AnERG, E. (1950). Barley and Wheat from the Saqqara pyramid in Egypt. Lantbr.HoDgsk. AmI.., 17,59-63. (Abstr. in BioI. Abstr., 25, No. 26373, 1951.)
ACTON, E. H. (r893). Changes in the reseIve materials of wheat on keeping. A'In. Bot.,
Lond., 7. 383--'7.
ADAMS, J. D., NIL\N, R. A., & GUI\'TH.'JUrr, H. M. (r9SS). After-effects of ionizing
radiation in Barlcy. 1: Modification by stonge of X-rayed seeds in oxygen and
nitrogen. Norlhw. Sf;;" 29. 101-8. (Abstr. in NudUlT Sd. Abstr., 9, No. 7632,

1955)
AFANl'SLEV, M. (19-J.3). Germinating Nandina damrstic/1 seeds. Anur. NUTJnym., 78(9),
5-6
AHLGKF.N, G. H., F!SK.E, J. G., & DouE.:'\"KO, A. (1950). Viabili[}' ofbromcgrass seed
a.~ affected by dehul1ing and by storage in fertilizer. Agrofl. J., 42, 336-].
AKAMINE. E. K. (I9.B). The effccr of temperarure :md humidity on viabilif)' of stored
seeds in Hawaii. Bull. HarIJaii Agric. Exp. Sta., 90, 23 pp.
- - (1951). Viability of Hawallan forest tree seeds in storage at various temperatures
and relative humidities. Pacif. Sci., ;, 36-46.
- - & RIPI'ERTON, J. C. (1938). Gcnnination and viability of crop seeds in Hawaii.
R~p. Hawaii Agric. Exp. Sta., 1937, 10-12.
AKI, S. & WJl.TA."<ABE, S. (r955). Studies on the storage of the seeds of Kintoki carrot.
I: Influence of storage temperature and seed moisture on tbe germination of
seeds. Kagawa KmrilS1l N6ka Daigaku, CakujulSu H6koku (Tuh. Bull. Kagawa
Agne. CoIl., Japtm), j(I), 31--:35. (In Japanese, with English summary.)
AUlO, G. (1g08). Les enzymes ct la faculte germinati\'e des graines. Bib!. Ulliu. Arch.
Sri. Phys. It Nat. Sa-., 4 25, 45-52.
AUL"J, G. S. (1957). Storage beha...i or of conifer seeds in sealed containers held at
o"F., J2"F., and room temperature. J. Fcr., 55, 278-81.
ALTSOil!L, A. M. (J949). Swcct and unspoiled. Chemical trcattnent prevents deterioration of cottonseed under storage. CIIlln. Ind. Phil., 64 (2), 2tl.
- - CoNlJON. :M. Z. & L ..MBOU, M. G. (1951). Method ofpreycotion of deterioration
in seeds. U.S. Pat., No. 2,571,01)5.
AMATO, F. D' (1954). Di ak-urn fisiologiei e genetici dell' invcchiamcnto dei semi.
Contributo al problema della st:nescenza e dell:t rnUlabilici spontanea nei vegetali.
[On some physiological and genetical aspects of seed ageing. A cOIllTibution 10
the problem of ageing and of spontaneous mutability in plants.] Caryologia, 6(2)
217-40. (In Italian only; abstr. in BioI. Ab!tr., 29, No. 29695, 1955.)
ANOF.RSON, J. A. & ALcoCK, A.. W. (Ed.) (195+). Storage of cereal grains and tIJdr
produr;ts. Monog. Sn. Aflln. Ass. C(ml! Cliefl/., wI. II, 515 pp.
M'DRf:N, F. (r945). LagringsfOrs6k med betat uts:ide. [Storage experiments l't-ith
disinfected seed.] Vri),'lskyJJs/lutiM'r" Srorkh. 1945 (I), 1---6. (In Swedisll only;
abstr. in Rro. Appl. /\1)'{01., 25. 207, 1946.)
-The citation of an abstract indicates that the original publication was not seen.

166

UIJ3LIOGRAPHY

ANON. (I8.f-3). Mummy wheat. Gdnrs'. ehron., 1843, 78~.

- - (1928). Noble fir seed keep well in cold storage. For. Wrkr., U.s., 4(4), 14--15.
- - (1931). Regulations for warehousemen storing seeds under the United States
warchousc act. U.S. Dep. Agr. Bur. Agric. Ecoll. S. (5 R.A., No. 122, pp. ZI.
- - (1932). Influence of age of pumpkin seed on yield. Qj. Agrie. J., 37, 74.
- - (1933). Samen aus mumien. Mill. DUck Dtmdrol. C(s., 45, 345.
- - (1934) Mummy wheat. Nature, Lalld., IJ4, 730.
- - (1935). Valuable information for sccd~men (Bodger Sews, Ltd., E1 Monte, Ca1.).
F!or. E.-reh., 8S(I), 5.
- - (1936). Connccricut Agriculturnl Experiment Station. Reparr of the director for
the year ending October 31,1935. Bull. eWIII. Agrie. Exp. Sto., 381, 165-20Z.
- - (1940). Air-condition seeds. Am(T. NursrT')'m., 72(5), 31-32.
- - (l940a). Cinchona seeds lose viability mpidly. Rep. P.R. Agrie. Exp. Sta., 1940,
18-19.
- - (19400). A satisfactory method of packing palm seed for distant transport.nion
was developed. Rep. P.R. Agrie. Exp. Sra., 1931)-40, 62-63.
- - (1941). Storage ofred pine seed. Tech. Notes Late St. For. E.TP. Sta., I p.
- - (I94IQ.). Storing coffee seed. Trop. Agriculture, Trill., 18(z}, 25.
- - (194Ib). Planes taking rubber seed from Canal Zone to Era:rlL Chf(m. BOI.,
6(q),3 2 5
- - (194U). Viability studies on tree and shrub seed. Rep. N.Y. St. Agric. Exp. Sta.,
1939-4, p. 4I.
- - (1942). Duration of viability :in seeds. Gdnrs'. CiTron., I I 1(2893), 23-1-. (Abstr. in
Bio!. Abslr., 17, No. 6934, 1943.)
- - (1945). Good bread made from twenty-year-<lld wheat. Sci. News Lm., Wash.
47,18..;..
- - (1947). Test seed~ by electricity. Am~r. NUTSnym. 85(1Z), 6.
(1948). Woody-plant seed manual. Misc. Pub!. U.S. Dep. Agric., 654, 290-2,
361 -3.
- - (1949). Dry cold storage of seed corn. U.S. Dep. Agric. Rep. Agric. Exp. Sta.,
1948,43-44
- - (1950). Seed com keeps best when stored near zero in tight, dry container.
Crops f5 Soils, ::.(9), 29.
- - (1951). Ancient seed.. not viable. Sci. News Lm., Wash., 59, 180.
- - (195 1Q). Deep freezing prolongs viability of sugarcane seed. U.S. D!p. Agric.
R(p. At/minis/rator of Agnc. Res., 1951,379.
- - (1952). Low tempemtu1c storage keeps sugarcanc seed viable IlliIny months.
Crops (s Soih, 4(8), 27.
- - (195za). Manualfor t!sling agricultural and l~g(toble suds. Agric. Handb. U.S.
Dep. Agric., No. 30, Washington, D.C., 440 pp.
- - (r953). Longevity of Halogeton seeds. Mon. Bull. Calif. Dep. Agric., 42(4), Zgo---I.
- - (1953a). Recent research on drying and storage of rough rice. Sn. Bul!. SO/lfherll
C()-Qp., 29. 29 pp.
- - (1954). Losses in agriculture. R!s. Ser. U.S. D(p. Agric., 20(1), 95-101.
- - (19540). The preservation of viability and vigor in vegetable seed. ASGROW
A1allagr.,2, 32 pp.
- - (1956). International rules fur seed testing. Proc. bl/. Sud Tw. Ass., 21, 1-80.
- - (1958). Ne,,"~ Briefs. Sci(l/{(, 128, 1563.
ANTI-lOl',y, K. R. M. & TARR, S. A. J. (1952). The causes of deterioration of cotton
seed in the equatorial province of the Anglo-Egyptian Sudan. EII/p. J. Exp.
Agric., 20(77), 56-65.
A&\.I, M. & KATAOKA, T. (1956). Ecological studies on AIopecurus atqualis Sobol.
0) Influence ofsoil moisrureon the donnancy and longevity ofseeds. (4) Seasonal

,67

,
(

SEW PRESERVATION Al"\'D LONGEVITY

variation in the viable seed population and its vertical distn"bution in the soil.
Prut. Grop Sci. Soc. Japan, 24(4), 319-2). [In Japanese, with English summ:uy,
P3 2 3]
Aru.IOJ.TK, N. & DICKSON, J. G. (1956). wlinimum humidity requirement for germin:l.hon ofconidi:l. offuugi associated with storage of grain. Phytopathology, 46, 462-5.
- - - - & DICXSON, A. D. (1956). Deterioration of barley in storage by microorganisms. Phyl()plJ/hology, 46, 457-61.
AR.'\,'DT, C. H. (11}46). The internal infection of conon-sttd and the loss of viability
in storage. Phylopal!lOlogy, 36, 30-37.
ATANASOFF, D. & ]m-lNSON, A. G. (1920). Treatment of cereal seeds by dry heat.
J. Agric. Res., 18,379-90.
ATKINS, W. R. G. (1909). The aboorption ofwatcr by seeds. Sri. PrOf. R. Dublin Sac.
[N.S.], 12, 35-46. (Absir. in Exp. Stu. Rrc., 21, 725-6, 11)09.)
AVtJl.Y, A. G. & BL\lillSLEE,. A. F. (1936). Visible mUTations from aged seeds (Abstract).
AnJl!r. Nat., 70, 36-37.
BAILEY, C. H. (H)17). The moisture content of heating wheat. J. Am". Sa~. Agron.,

9.2.4S-5 T
- - (19170). The handling and storage of spring wheat. J. Amer. Soc. Agron., 9.
275-8 I.
B,\Il.LY, S. W. (r955). Air-tight storage of grain; its effects on insect pests. I: Calandra
granarifJ L. (Coleoplera, Cureulionidae). AIISI. J. Agri~. Res., 6(r), 33-5 r.
- - (1956). Air-right ston[re of grain; its effects on insect pests. II: Calandra ory;;ge
(small strain). Aust. J. Agrie. Res., 7(i), ri9.
BAIRD, r. D., i\1A0\1.~SfER. M. M. & RIST, C. E. (1950). Studies on a npid test far
the viability of com for industrial use. Cereal Chern., 27, 508-13.
BAKER, D., NEUSTADT, M. H. & ZElh'.'T, L. (T957). Application of the fat acidity test
as an index of grain deterioration. Cereal Chm/., 34> 226-33.
B.'LKKE, A. L. & NOF.a::ER, N. L. (1933). The relation of moisture to respiration and
heating in stored oats. Bufl. 10. Agric. Exp. Sta . 165, 31 / 36.
BALDwm, H. 1. (T93-1-). Germination ofrcd spruce. Plant Pilysiol., 9> 49T-532.
- - (r935) Catala:;e activity as a measure of viability of tree seeds. Am". J. BI)/.,
22,635-44
- - (1942). Fortst trt!( sud of tht north tell/perote regiolls witil special rtftrl!1lu to
Nortil AII/eriea. ChroniQl Botanica, Waltham, 1-lass., U.S.A., 240 pp.
- - & PI..E.\fJON, F. (19-1-T). Rapid method for testing white pine germination. Po).'
For. Nota, No. 30.
BALl., G. J. ('935). Serious losses in germination of seetl Qln generally be avoided.
Plor. Rro., 75(1937). ur-20, 49-5T.
llAR,\cR, H. & D.USKOY, F. (1955). Notes on stonge of Douglas fir seed. Proe. Int.
Sud Test. All., 20, 57-6i.
BARRONS, K. C. & Ald.UN, D. M. (19-1-5). A study of the causes oflow gcnnination
of radish seed crops. Quart. Bull. Milh. Agnc. Exp. Sta., 27, 3g8--1-08.
BARTLETT, L. (1859). Vitality of seeds. Report (if the Commissioner o[ Patents [or Ihe
Year 1858, Washing/on, D.C., pp. ]]2-].
BARTON. L. V. (1930). Hastening the germination of some conifcrou... seeds. Allier.
J. Bot., 17, 88-II5. (Contr. Bo)'({! Thompson Inst., 2, ]15-42, 1930.)
- - (1932). .Effect of storage on the vitality of delphinium seeds. COnlr. Boyu
Thompson hISf., 4. LP-54-.
- - (r933) Germination and storage- of ,delphinium seed. HI/ll. Aml!T. Delphil/ium
Soc., 2(1); 12-14. (Prof Pap. Bo)'u Thompson Insl., 1(26),2-1-8-50, 1933).
- - (1935) Storage of vegetable. seed~. COlllr. Boyce Thompson Inst., 7, 323-32.
- - (19350). Storage of some coniferous seeds. COnlr. BOJ'u Thompson hw., 7.
379-404.

:
:

mnLlOGRAPHY

L. V. (lIB9). A further reporr on the storage of vegetable seeds. COllir. Boyce

Thompson Insl., 10, 205-20.
- - (19390). Storage of elm seeds. Confr. Boyu Thompson Imt., 10, 221-33.
- - (!93Qb). Storage of some flower seeds. CIJTlIr. Boyce ThompwlI !1m., 10, 399-427.
- - (1939'). Germination and storage ofWy seeds. Nat. Hort. Mag., 18, '93-4.
BARTON,

--,.

- - (193gd). Experiments at Boyce Thompson lnsriwre on germination and domlaney in seeds. Sci. HQrl., 7, 186-93.
- - (1941). Relation of Ccrt::lln air temperatures and humidities to viabilily of seeds.
Contr. Boyce TlwmpsonIlISt., 12, 85-102.
- - (1943). Effect of moisture fluctuations on the viabilily of seeds in storage. Ccm/r.
Bo)'ce Thompson bISt., 13. 35-46.
- - (19430). The storage of cilrus seeds. COlllr. Boyce Thompson but., 13, 47-55.
- - (19-1-5). A note on the \;ability of seeds of lvlag:I, Montezuma spuiosissitllil. COnlr.
Boyu Thompson Inst., 13, P3--6.
- - (19-450). Viability of St:eds of Frll.,iIllIS after storage. COlllr. BGyu ThGmpSGl1 11Ist.,
13,427-32.
- - (1945b). Respiration and germination studies of seeds in moist storage. Ann.
N.Y. Acod. Sri., 46, 185---208.
- - (19-1-7). Effect of different storage conditions on the gl-~mination of seeds of
Cindwlla Ledgt:rian~ r-.1oens. Contr. BG)'u Thompson Inst., 15, 1-10.
- - (1948). SlOrage of seeds of the Rl-'gal I.ily. Boyu TJJOmpstJl/ !lw. Prof Poper,
z(6},45-51.
- - (1949). Seed packers and onion seed viabilil). COlltr. Boyu Tho/llpjOll IlISt., 15,
3-1-1-5 2.
- - (1953). Dormancy in seeds. Rep. I3th Int. Hart. Crmgr., Lond., 1952, 2, 1001-I:Z.
- - (19530). Seed storage and viability. COl/fT. Boyce ThompSO/l IIIS1., 17, 87-103.
- - (1954). Effect of subfreezing temperatures on viability uf L..mift:r seeds in storage.

Comr. Boyce T!lompson blU., I8,2J---'24.

- - (19544) Storage and packeting of seeds of Douglas fir and Western hemlock:
ClJTltr. Boyu Thompsrm Inri., 18, 25-37.
- - & CR(X;I,.'"l:R, w. (J948)~ Twenty years ofseed reuoreh 01 BoJ'u 1110lllpson Imtitute
for Plant Rtstorch. Faber & Faber, Londoll, 148 pp.
- - & GARMAl\', H. R. (19-1-6). Effect of age and storage condition of seeds on the
yields of certain plants. COlllr. Boyce Thompsoll IIlSt., 14, ~3-55.
- - & HOT01K..ISS, J. E. (1951). Germinatioll of seeds of Eidlhornio crossipes Solms.
CoMr. Boyu Thompson I nst., 16, 215-:lO.
BARTON-WRIGHT-sa WRIGHT, E. C. BARTOl"-.
BATF.'i, C. G. (1930). One-year storage ofwhirc pine sced.]. For., 28,571-2.
DATTU:, W. R. (1l}48). Effect of scarification on longevity of alfalfa seed. J. Amer.
Soc. Agron., 40, 758---9.
BAYFlllD, E. G. & OT)Ql'''''E1.J., W. W. (1945). Observations on thiamin content of
stored wh~t. FoodRes., 10 (6), 485-8. (Absrr. in Biol.Abstr., 20,No. 7183, 1946.)
Bf_~TI"IE, J. H. (1931). Peanut seed may be kept for several)"CatS under proper conditions. Yearb. Agric. U.S. Dtp. AgTic., 193J, 4-Z6-?
- - & BoswF.u.., V. R. (1938). Longe\;!}' of onion seed in relation to storage conditions. Proc. AIIICT. Soc. Hart. Sci., 35(1937), 553.
- - - - (1939). Longevity of onion seed in relation to storage conditions. Agri,.
Cire. U.S. Dep., 512, 23 pp.
--JACKSON, A. M. & ClJ"RRIN, R. E. (1932). Effect of cold storJge and age of seed
on germination and yield of peanuts. Agric. Cire. U.S. Dtp., 233, 12 pp.
__ & KlJSI-L\I..-\..N, L. J. (Ig.!?). Effect of moisture contents of peanuts on shelling
damage. Peolluf J. 011I1 Nut World, 27(1), 35. (Abm. in Biol. Abstr., 23, No.
I643I,lg49)

SEID PRJ::SERV.J,TION

.J,.".m

LONG.VrlY

J. H. & TAnlA."l, E. C. (1950)' Long--tenn stonge of parsnip seed. Proe.

Amu. SO(. Bon. Sci., 55, 435-'].
~ P. (I~). Sur b penn6J:bilite aux g:tt de I'atmosphere, du ICgumnlI de
certaines gmm:s llcssCcbecs.. C.R. Acad. Sci., Poris, 138, 1347"9.
- - (1907). Recherches sur Ia \;e Ialemc des graines. Ann. Sci.1\'ot. 9 so. Bot., 5-6,
BEATTIE,

193-3 I I .
- - (1934). La longcvite des graines macrobiotiqucs tr:Insmise par Louis Mangin.

C.R. Arod. Sci., Paris., 1990 1662;Bt:."....rrr, N. & Loo.\us, W. E. (HH9). Terrzrolium chloride as a test reagenl for
frt:czing injury of seed Com. P14nJ Ph)'sioI., Z.J, 162-74.
Br.R."l,u., D. M. (1953-54). The use of 2, 3, 5-triphenyltctr.lZOlium chloride for the
dctcrmin:uion of the viability of seeds. Colomb;", Mi"iSl~,ja Agrje., Dit,. InI'(st.,
Infonn. Tmtua, 1,79-121). (Abslr. in CMm. Ahstr., 49, No. 1178Z, 1955)
BIBBY, K. ~'L & WlllJA.\lS, R. W. M. (1953). The effect of
o\"O"-",;mO" storage
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Ifl

-,

SI::J:D

P~V.-\TION

_"-'ID

LONGEVITY

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429""44 (In Danish, wilh English summa.ry, pp. 443-4.)

(t948). Germination of buried ant! dry stored seed. II: 1934-+!-. PrOf. Int.
Sud Tfst. Ass., 14, 19-26.
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in ruhcnden Samen. Bd. Bal. Zbl., 48, 22l}-3I3.
K.WDSON, L. (1941). Investigations on dchyed photOSYIlth~ in chlorophyll_bearing
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Agn"e. Exp. SIll., I~J40, 117.
KOF.HiDt, B. (1938). Effect of prolonged storage of treated seed com (Abstract).
PltytfJpatho!og)', 28, 13.
-

,
,-

BffiLIOGRAPHY

KOEHLER, D. & BE'i'F.R, W. M. (r956). Effect of fungicide and of storage temperature

on fungicide inilll"Y to wheat set:d. Plum Djj. Rep'r., 40, 490-2.
KON"DO, M. (1926). tiber die Da...!lef der Erhaltung dec Keimkraft bei verscbiedenen
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~

a granary and Ute hulled rice stored therein by a desiccating materiaL Ba.

Oltara Insl., 7. 227-'37_

- - & KA.<;AHARIr., Y. (1941). [A repofl on storage of seed.~ for JO yrs. wilJl calcium
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No. 10581, 1950.)
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chloride, with special reference to the underdried product. I. Ber. Ohara Imt.,

6,335-9
- - - - (1935). Storage of rice. XUI; Storage of rice ifl tin containers wirh calcium
chloride, with special reference to the underdried product. II. Ber. Ohara Inst.,
7,9g-10Z.
- - - - (1937). Storage of rice. XV J: Storage of rice in concrete silos for five years.
Her. Ohara Imt., 7, 47J-SI.
- - - - (1938). Storage of rice. XV'III; Rehrion between varying moisnue content
and ehange in_quality of hulled rice, with special reference to the underdried
product. Ber. Olwra Imt., 8, JJ-3-l.
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hundred years in a granary. Ba. Ohara Imt., 8, 47-52.
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moval of moisture from hulled rice by heated air. Ber. Ohara Inst., 8,
35-46 .
- - - - - - (I94-1). [Oll the storage of seeds of forest trees. I.] Amer. Sci., 32,
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- - & TERASAKA, Y. (1936). StorAge of rice. X\l; C.ompan'iOn of calcium oxide and
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Ohara Inst., 7, 32g-34.

ISr

SE!ill PRESERVATION k"\'D LONGEVITY

v. M., Z'UlEt.INA, Z. K. & SHOif:RBAKo\', V. G. (1956). Influence of

moisture and some of the biochemical factors ou the conditions of storage for
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KORNFELD, A.. (1930). Unrerwchung dcr KcimIahigkeit mit Hilfc \"on Farbl6sungen.
FOrluhr. undw., 5{20), 682-3.
KRF.ITLOW, K. W. & G!JUlR, R. J. (1946). Viability of stored seeds of forage crops
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KRETOvIOI, V. L. (1945). Physiologico-biochemical bases fur the storage of grain.
Acad. Sci. S.S.S.R., [nsf. Biochl:1ll.,ltJoscow-uningrad, 136 pp. (Abstr. in Chl:1ll.
Abstr., 40, 1237-8, 1946.)
- - & STARODUBTSEVA, A. 1. (1956). Respiration and lipolytic activity of stored
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KRL'YER, T. (1939). [The storage of seeds of the quinine tree.] Sovmlt. Suhtrop.,
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Ktn\LE, R. E. (1882). The germination and ~itality of seeds. Torr~y Bot. C/. S~porof~.
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KUROKAMI, T., EHIHARA, T. & TAKF_\UTSIJ, T. (HJ47). Studies on the development
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KOPE!KOVSKli,

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'9.

183

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,,-,,1)

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c.

(Abstract.) PhytOpalhology. 31, 768--9.

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'9 '

SEED PRE.'lERV,\TION A1\'D LOKGEVln"

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Trop. Agrie-ultIlTt, Tnn., 24, 51-53.
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- - & VlU.AF.<\Nf.., A. G. (1947). Seed storagc. Rep. P.R. Agru. Exp. Sta., T946,35.
WmIT, J. (1909) The fcrments and latent !:ife of resting seeds. Proc. Roy. Soc., B. 81,
41?-42
WHITE, O. A. (19-1-6). The germ:ination of pe:IS in Florida and King Tut's tomb.
Turrox News, 24(1), 3 pp.
WHYMPER, R. & BR.IDI..E't", A. (1934). Studies on the vitality of wheat. I: ProlOn.,":ItiOll
of vitility of wheat seeds. Cereal Clttm., I.l, J4,9-60.
- - - - (1934<'). Studies on the \itality ofwhcat. II: Influence of moisture in wheat
seeds upon imbibition and speed of germination. Cereal Chm:., II, 546-50.
- - - - (19341. Studies on the vitality of wheat. III: Vitality, anothe action of
heat on wheat seeds. Cereal C!lem., II, 625-36.
- - - - (1947) A note on the viability ofwhe:It seeds. Ct:TeIJI Chelll., 24, 228-g.

'96

-;;
-

nIULIOGRAPllY

\VIlLIAl-1S, M. (1938). The moisture content of grass seed in relation to drying md

storing. IVeIlhJ. Agric., LJ, 213-32.
Wll.SON, A. K. (1953). Stor~gc and germination of Douglas-fir sttd in central Idaho.
Res. lv"ous, Intermountain FOTU! {5 Rallgt Exp. Sto., Ogdm, Utah, NO.5, 3
Mimco pp.
WINTERS, H. F. & RODRIGUU-CoJ.ON, F. (1953). Stol'llgc of Mangostccn seed. Proc.
Amer. Soc. Hart. Sci., 61, 14---6.
WOllF1'o'wEBrn, H. W. ([942). Ubcr die Lebensdauer yon Kurtolfe1=en. Angell',
Bot., 24(2), 259-60. (Abstr. in BioI. Abslr., 23. No. 16854, 1949.)
WRlGIIT, E. C. BAKTO:".'-,lloom, R. G. & PRiNGLE, W. J. S. (19..1_1.).Anal}'sis of barley
from King TUlanlhumcn's tomb. NatuTI:, !.Amd., 153, 288.
WRIGHT, R. C. (1941). Investigations on the storage of nuts. Tuh. Bull. U.S. Dcp.
Agri(., 770, 35 pp.
WYTffi."ffiAOl, E. (1955). Dcr Einfluss vcrschicdcncr Lagcrungsfaktorcn auf die
Haltbarkcir von Fddtimcrcicn (Luzerne, Rotklce, und gemcinem Schotenklec)
bei l:ingcr daucrmlcr Aufbewahrung. Landw. Jb. Scltweiz, 4. 161-(}6.
YA.1'''AGISAWA, T. & ASAKAWA, S. (1953). 'Woody plant seed viabilil)' tests with tctrazolium salts. J. lap. For. Soc., 35(2), 4<>-42. (Absrr. in BioI. Abslr., 27, No.
23-1-62, 1953)
ZAO-IARIEW, B. J. (1939). Em bcqucmcs SchnclIke.imvcrfahrcn zur Priifung des 5:unens
einiger Naddholzartcn. Forsrwiu. Zhl., 6I(8), 238-49.
ZtJ...,''Y, L. & COLL'L...."', D. A. (1938). TIIC chcmical dctcrmination of soundness in
corn. Tuh. Bull. U.S. Dtp. Agric., 6+4, 23 pp.

,
:

CONVERSION FACTORS OF WEIGHTS AND MEASURES

Mr'Uura oIlY.itM-AI;\')il1u!><>is /0 Mtlri<:
I

dram (dr.)

I
,
I
,
I
I

Cum", (0:<")
pound (Ib)
.lolle [.1.)
quart",. (qr.)
hllo<!n'<l_..gbt (0"".)
(long) too (2,240 lh)

=7'~J
15

1'77' II"'''''' ~ )

sraio..

drams

0"4,.l<iltIgr<o"..

14 POUIlW

6'350 kilorams

, stones

"?Ol kilogra"",
;'10-80: \<llot,alIK
"o,6lnnnes

4 quart=
=0 bundred,.'eight
J1d,~

U> Aroi,Jupeif

, milJigJ;l.m ("".j
I gram ((m.l
1

kiJosram

&""'"

'8'3

'60uo",,"

gnuo
o'sl4 dram

O<{l'S

r,ooognma

(kI:.)

, IOOne (melric Ion)

2'20;'1 p<lullds;

',<>< kilognm<

ton

1l-gll4

A ",,,;co,, II'..~hJ. la Mark

, pound
I ~n!21

4:13';'19' gnrns
4"3:19 kU<ogr.>=
tonne

'00 poun'"

, (<hert) Ion (:,000 lb)

20

<>ental>

O"'}OJ

Mdrioto A",,,,,ie<l..
100 kilog,-""", (kg.)
I tonne

W<i~AlJ

I quintal
1,000 kilogr:l.ms

2':05
'"I\>'

cent"..
(5hortll"...

fJr""u.uDj L"'ll},-B,i,i.1t /0 Mel,;.

incb (in.)
I {oot (ft.)
I yard (yd.)
I mile
J

"5'4"" milliulet~

" iucheo

30'480

3 f..,t

ce,,([met...,.

""'lI 4 '''''Ins

1.76<> ;'Uds

'-60<;1 kilometres

M <IT'" /0 /lriti>!
I

mi"""n

tI'l

'1',000 mm..

'1'~,400

',""" m.

0'''39 incl,
0"394 il\Co
3"937 iocbeo
'"<>94 yaros
3'8' I_t
39"3]<1 ineoo
~6.. miJe

(l/I,06<>.Oo<> m.)

milli"""re I""".)
, <:entime,,.,, (em.)
d.nm.lre (dm)

me',." (m.l

, kilomet (km.)

M ....",-.; "I A."" (Ba.uJ ..a I

BnlWl 1.0 M

'quare inoo I~q, In.)

I .qnue tool s.q.IL)

'0,000

6.40

.qn"", millime,,,,

6'4~a ''1. o:ntim.=

""093 ''1_ metre
0"836 ''1. mel'"
"'40~ h""I.",
"3<)<1 kilometres
'36'998 httla=l

9 ''1, II.

..=
, .qure mile
1

39'37<> 'nelw)

ff4d.. _

d,,,,

H"l.in.

, 'qua" f;ud (.'1' yd.)

io<o

.q.m.

a<:reo

1.'1. mm.l

"qu.", ren,i""'I'" (''I. c"'-l

.quare. metf. {'q. "'-I
heel"'" (h,l
.qu= kilometre (sq_ km.l
Jt....

'00"'1. mm.
[00 "I. 11m.
'opoo sq. Ill.
'00 h...

"'a 01 Vol"....~B,iUd, 1.0 M,!>",

, cubic iocb (cu. in.)

cubic foot (cu. ft,)
, cuWc yard (en. yd.)

07 CII. I,.

eubi<: <=tti""'rn Icc, - ml.)

CIIbie decimetfe (cu. ilin.)
, cubic metre (cu. lD-1

cu, em.
,,00<> cu. dm,

',7=~C",

1~,3g7 ell. """timetres

=8'3<7 ctI, <tecl",e'fh

in.

""7~,

cu..

,,,,,In

,,'06, cu. inch

0"<>33 cu.. loot
f3ell cu. yam.

1,000

M .... ,,"~ 01 Cap...;ly --(lJ<oud on r",pnul! :allon _ 4'S 16 lil,a).

I pl"t {pt.J------<>-368 W[.

I talkm (s-al.}-r:H61iIns
pints_I quart {qL}

r )\mori""n !>,,1I0n-378, lit",

'Iitre----f,OOOcc.
4 '1U3.rls-, g-'l.ll"n

Mr.';e 10
pint
J

gaD"..

"'S~8

lit'"
4,~6Iit....

BTili.lt.

millilitre (mL _ =)-<>'06. cu. in.

I centililnl (d)-Io m1.-----<>6,o .ll. in.
, lit", ILI-."" cl.~'76<> pinU

Ttmpetalu

o Centil;r.Jde (_Celliur.)_3' Fab",ebtit

The 1"llowint l"rt""I~e amn...,' lh. t ..... =j"r thermometric =1",:
F. b",,,heit to c.-"ligrade; C - 3/<l (oy - 3')
Centigrade to Fahrenlleit: OF _ "Is ('e + 3')

19 8

SUBJECT INDEX'
Abdmosrhus urn/tn/us seeds, vi1bility detennincd by tcrrazolium salts and excised
embryo technique comp;=d, 128
Abies a/ita seed~, viability tested by U!;e of
indigo-carminc, 117
- baJsamra seois, romparison of germination,
e:<cised embryo, and tetrazolium TCS15, 130
(oncofor seeds, ,"iabiliry test by excised
embryo reclmique, lf~
- nobilis seeds, longevity or, 79
AbU/iton tkeophril51i seeds, life-span in soil, 9
-

Au,. =dIlITum, lc:lf, double =

:md seeds

of, Fig. I
seeds, effect of de!<iccatioll on viability of,
9-l--lJS
- taillrirom seeds, viability uetcnninoo by
te01lwlium salLS ,,"d excised embryo tech.
niquc compared, :l28

ACQffiS, rrr Qu",ctl5

AumlS calamus !iUds, longe,it), of, 92
Auinro ado,ala seeds, life-span in soil, 10
Agricultural seed~, \iability tcsteO with
tel:r:IzOlium salts,

zIi---7

Agrosltmrr.a gilbago seeds, life-span in soil, 8

Agropyron seeds, effcct of Dl:Ilurity on viability and longevif)' of, 38
AgroSlis !.tokilliftra seed.~, longevity of, 68
Air, elkct on longevity ofGossypium seeds, 34
- elfect on longevity of Ory~ satiw seeds,
35
- effect all longevity of Populus seeds, 3S
-elfcet On longe~ityofZta mays seeds, J.j.-J5
A/bi -~ill julibrWiIl seeds, Iife-sp.m of, 3
Akurius seeds, effect of desiccation on
viability of, 98
- - \;:l.bilit)" tesled by use of indigo-car~
mine, 117
Alf:l.lfa, su Mdicilgo wlioo
Alis",," plill/iago-aquillicil seeds, longe\'ity of,
91,9 2
Alkaloid content, of deteriot"<l.ting seeds, J 52
AlliulII ujJto seeds, absorption of moisture by,
I5--'Zt, :26
- - - effect of different packets on \'ia~
bilityof, JOI--6
_ _ - effect of fIlIlisrure on viability of,
14--21
- - - effect of rebtin' humidity on \'ia~
bilit}" of, 18, 25--28
- - - effect of tempet"<l.rure on viability of,

.S

- - effect of temperature and humidity

on S[Qt"<I.ge of, 1.;8

Allium rtpa seed., longnity in storage, 50-52

-

- -

"-

~fc

moisture content for storage of,

use of old, 141

Alopuurus a(quafis seeds, life-span in soil, 10

- frrlllttlsis seeds, longevity of, 68
Aisi/It mtJia seed., life-span in soil, 8
Alumaria, longevit}, of, on seeds, 45-.;6
Amaral/thuJ rrlrojlrSlu seeds, life-span in soil,
_

7,8

- - respinltion of, 12-13

- - viability on moist glas,;~wool ~I

20~C"

n-IJ

seeds, carJbse acti\;t}' as measure of

viabilitJ'of,II5
Ambrosia arum"iifolia seeds, life-span in
soil, 9
- rlalior seeds, life-span in soil, 7, 8
Amyl:lse activit} in rice se.eds and acorns, l51
Andropogt1!l seeds, longe\'lty of, 69
AI/gtli,a archangrU,a seeds, longevity of, 76
Angiosperms, vii
AI/themis ,otula seeds, life-span in soil, 8
Anrhqxlll/lhum odorarum seeds, longevity of,
68

AnthJ</lis vull/ffarill seeds, life-span of, 3

AI/tirrhinum majlls seeds, <:frect of belowfreezing tempenIture on, 33, 62
- - - longe"it), in SlOnIge, 6l
- - - mutltion r:lte affected by age of,
IH, 156
Apium grar;eo/ms seeds, s:Jfc moisture content for storagl.' of, 21
,apricot, Sf' Prill/US
Aquatic plants, dormancr of, 91
- -longe.vil)" of, 91-94
- - su also names of indh'idllal plants
AriUhis hypoga.ea sel.o;, ahsorption of moisture by, 15--21
- - - ehanges in proteins during storage of,
150
- - - effect of age of seed on plant quality,
138--9
- - - effect of moisrurc on viabilicy of,
14-21
_ _ _ effcrt of relative humidity on viability of, 18
- - - effect of temper:tture on \'lability of,

.,

- - -longe\'ll)' of, 74,5

- - - s:ttc moisture Wlllent for storage of,

"

Arnica nllmlllnlJ seeds, longevitr of, 76

.. M in the gener:tl ten of this book, no distinction is here made between seeds and ~eed
lil.:c fruits, all being commonly referred to as 'sceds',-Ed,

'99

SEED PRE.')ERVATION Al\'D LONGEVITY

Ascorbic: add in Vign.. s';n,nm, destruction
ouring stornge, 152

AJlh,

se~ FraJ."inu~

Asparagus plumams seeds, effcct of bclow~

freezing It:mper:tllirC on, 33, 6::
.I~p(:rgillus amdidus, effel;{ of wheat grain
inY;l.sion by, 41, 43
-flaws, effect of whC:It grain invasion by,
- - effect on deterioI1ltion of l;r.lrley seeds,
Plate 3
- nig.r, cffect on deterioration of barley
seeds, Plate 3
- repms, effect of ..-bC:It grain im"asion by,

on

4 1,43
restriclUs, effect of wheat grain invasion by,
-10, -I', 43, Plate 2, Fig:" 7
Aster, Chirut, su Caflistephw chinensis
Astraga/us l1lassilimsis seeds, life-span of, 2
- utrig.r seeds, life-span of, ]
Aubergine, su Solar-urn mdongma
Austrnlian pea, See Dolichas lignams
Auxin content, of deteriorating seeds, '52
AZ,'e1Ul datiar seeds, longe,if]" of, 68
- sali'!.YJ seeds, tetrazolium lest for 'iability
of, '2', a3, Fig. 12
A:ronaplls afjinis seeds, longe'if]" of, 6g
-

Bacteria, cff~"Cl: on longevity of seeds, -10

- stained by tetr:lZolium, 121, 122
Bamboo, See PhylllJSlachys
Bark:y, see Hordeum vuJgau
- seed.~, Egyptian mummy, \':iability of, 5
Bean, su Phasealus
Beech, su Fagm
Beer, su Beta nulgaris
Begonia seeds, longe~ity in storage, 6,
B.rb(ris thunbagij seeds, viability determined
by letr.lwlium salts :md excised embryo
technique compared, 128
- - - \':iability lest by excised embr}"o
technique, 113
Jhta vulgaris seeds, effect of age of, on plant
qualit~r, t40

- - - effect of rebrive humidity on ,';abil~

ityof, 18
- - - effect of tempcnture on viability of,
-

"-

- longevity of, 75--"]6

- - safe moisture content for stonge of,

"

Belulll seeds, longevity of, 87

Bibliography, 166--q7
Degnania (C.:mpsis) rtJJicans seeds, viability
test by excised cmbI}"O technique, II;;
Biochemical tests for seed viability, report,
135--6
Bismark brown, for resting seed \iability, uS
Blue lupine, su Lupinus I,irstllus
Birch, see Baula
Blueberry, sce Vaccinium
Brassica nigrlJ seed.~, life-span in soil, 7, 8
Bra~sica seeds, cifect of age on plant quality,

'39

- elf....;t of light on keeping quality of, 36

- effect of rebti'-e humiditv on viabilih'
of, ,8
""

Brllssica seeds, effect of rempernure on

bilityof, IS

yia~

- - 10ngl:\'It}' in storage, 5O--P

-

- safe moisrure coment for stora;::e of,

"

- - staining of, Iz6

Dromegrass, see Bronzus
BrOIll~ ina",is seeds, effcct of Jrulrurity on
,;ability and longc\';f]- of, ;;8
- marginalus seeds, effect of m::mrriry on
viability and longe,"if]" of, ]8
- palJanthus suds, cltea of maturity on
viability and longevif]' of, 38
- ucaNnus seeds, life-'iipan in soil, 8
- seeds, effect of mechanical dehulling On
longevity of, 37
Boried seeds, Ue Seeds, boried in soil
lJursa bursa-piIStoris seeds, life-span in .'>Oil, 8
Bu/amus umbel/aws seeds, 10n;!C,';ty of, 91""""92

Cabbage, sce BrassiclJ

seeds, effect of desiccation On ,iabilit)'
of,9 8
Cajanus indicus seeds, cata1:rse al"tivity liS
measure of 'iIbilltJ of, 116
Calla paIus/ris seeds, longeyity of, 92
ClJlIicarplJ jJUrpurea sel:ds, viability test by
creised embf]-o tcclmiquo; 114
Callistephus chinensis seeds, drect of age on
planr quality, 141, 1-\2-3
- - - effect ofbelow~freeling temperatures
on, 62
- - - effect of oxygen and p:miaI ,,,,cuum
on, 35
- - - longe'-lry in storage, 56~57
Calli/rich, aquatica seeds, longevity of, orCampsjs radicans ~ds, ,';ability determined
by tetrazolium salts and excised embryo
technique compared, uS
- - - su illS<} Bignllnia radicans
Canna plJnicullltil seeds, life-span of, 3
Cannllbis sat;vll seeds, Ylability tcst by ex~
clsed embryo lI:chniqne, II-I
UnnaCl':::l.e seeds, life-span of, 3
CapsdIa, see Bursa !mru-pasloris
Capsicum frul~suns seed5, effect of age on
plant quality, qt, l43-4, PI. II
- - -longevity in stora;:-e, 51-52
Caragllnll arkrnrens seeds, ,iability tested
by lISe of indigo--carmine, 1'7
(".arbon dioxide, effect on longe\it)" of
CosUpiu"" seeds, 34
- - effect on longe'"if]" of Oryza JilliiJIJ.
seeds, 35
- - effect on longevity of 7.,a mays seeds,
CIUlW

34-3-

Carpin~s seeds,

rules for testing- with tetr.!zolium salts, 135

Carrot, su Dazu:us carOlil
Caryll illinomsis seeds., Iongenry of, 89
Cassia bicapsulilris seeds, life-span of, 2
Cllssia ,""lIijU.ffil seeds, life-l>'J'lIn of, 2
Castor oil, su Ricinus communis
Catalase activity, me:lS\lrement for testing
~eed \;ahilif]', Il5-11, '5'

'00

SUBJEcr ll'.'DEX
Calalpa !puio!1l seeds, viability det~TTllined by

tetra7.oliurn salts and excised embryo !echniquc compared, ~28

Call/ryll seeds, longevity in storage, 61
Cauliflo....cr, S<!~ Brallicu.
Cedar, wes!ern red, sa Thuja pli&ara
C~/a!rrus sramlms seeds, viability determined
bv LClnIzolium ults and excisl::d embryo
technique comp:rred, 128
_ - - viability test by excised embryo
technique, II]
Celery, sa Apium grll~'eolern
Cmtaurea raldtrapa s~ds, lif<:-5pan insoil, 10
_ r~pe"s seeds, lif<:-5pan in soil, 10
- sol.titialis seeds, life-span in soil, 10
Crphllrosporiu", /urrmonium, longevity of, on
se<:d., 46
Ccratllph)'llum dOl/asu", seeds, longcvicy of,
9'
Cnds ca'lad~n5isseeds, yiability determined by
tetr;lzolium salts and acised embf)'O technique compared, 128
_ _ - viabilit~ Wit hy excised embr}'o
teehniqu", II]
CaclIspura Imirola, longevicy of, on seeds, 46
Cereal seeds, effect of rcLuive humidity on
moisture content of, 18
- - longevity of, 67
_ - Y'iability tested with tetrazolium $:tIts,
IU--6, Fig. 12
- - sa also names of individlll1l species
Chamomdcs japlInira seed~, comparison of
exei5cd embryo and tetrazoliumtcstsof, 132
_ - ~ ,'iabllity test by excised embryo
technique, 1I4
Chactorhloa /uresccllS seed~, life-~lUn in ~oil, 8
Chamaesyrc marulata scrds,life-span in soil, 8
Chemical composition, of det~noratingseeds,

Il-50

- treatment nf seeds, for control of fungt or

insects, 46-49
- - - - for prevention of deterioration, 48
- - - ~ for resting seed viability, IIJ-20
Cherry, Ut Prrmus
Chestnut seeds, transportation of, T58'"9
Chewing's fescue, Se! Ptstura ruhra
Chi~nanJhus virg;nira seeds, development of
embryos on moist filter_~per, PI. R
- - - viability determined by tetrazolium
salts and e~ei.~ed embryo technique compared, Izg
- - viability test by excised embryo
technique, 114
Chokccherrv, sa Primus
Chr)'Sanrr.cmum spulabi!! seeds, effect of
below-free-.cing temperatures on, 62
Ciur ariclum seeds, cab.b;;c activity as mC<lSurc of 'l'iability of, 116
Cinchona lragmana seeds, effect Qf light
during storage on, 36
- - - 10ngC'';t} of, 8g--Qo
Cirrul!us ntlgaris seres, safe moistureeontent
for storage of, 21
_ _ _ n.abiliry test hy c,"tcised embryo
=hnique, II)
Citrus seeds, effect of desiccation on viability
of, 96-97

Citrus seeds, selenite (cst of, 119

Gover, su Trifolium

Cocklebur seeds, tetrazolium (cst of, 1Z6

Coffia seeil$, effect of desiccation on viability

of,98
Coltsfoot, su Tussi/ago farfara
Congo red, for testing seed viability, 1I8
Conifer seeds, effect of below-frCC7ing storage on, 33
- - effect of tempcrarore on viabilicy of, 30
- - longcvity of, i!l-a)
- - tetrazolium testiug of, 1)0-1, t33
- - X-ray testing of, 137
_ - su also llilmes of indi\'idual species
emmofrufus upium seeds, lifc-lip:m in soil, 9
Coreopsis seeds, viability lesl by ~~dse"
embryo technique, II]
Coriandrum salitoum secds, ,';abilicy in stora.,re,
5

Com, StC Z~a mays

Comus seeds, ,';abilicy determined by tetrazolium ~':Ilts and e~cised embl)'o technique
compared, uS
- -:- ,'iabilicy test by excised cmbryo tcrh",que, II)
Cotton, sa Gossypium
Cowpea seeds, )';clds of old and young, 139
Cr(ltae!fll' rTuf-ga/li seeds, deve1('pment of
cmbI)'o on moist liIter-raper, PI. 8
Cralatgus spp. seed~, viahllity test by excised
embryo technique, 114
Crcpis seed~, mutation ra!e affected by age of
seeds,153--{i
Cresol red, for lesting seed viability, (18
Cucumber, see Cucumis
Cucumis _1o seed~, \'iability in storage, 50
- sati,,'Us seeds, viabilicy in storage, 50
- seeds, effectohge ofseed on plan! quality,

",

- - effect of moisture on 'viability of, PI. 7

~ - !;:Ife moisture content for storage of, 21
- -:- \ubilicy test by excised embl)'o techroque, II)
Cururhira jl!PO seeds, yiability in storage, 50
- seeds, effect of age on plant quality, 138
- - effect of fmit nJalurirv on keeping
.
quality of, 39
- - viability !esc hy excised embl)'o technique, II)
Cllpllliferae se<:~ effect of desiccation on
viability of, 98
Cydonia japonita, sa Chamomdes japmlicll
ohllfflga seeds, eompanson of excised

embryo and terrazolium tests of, IJ2

CJ'nog!ossum amahi/e seeds, "iahility test b)'
ex~-ised

embryo technique, 113

CynosUTlls "is/a/us seeds, longe\'il)' of, 68

Cyrisus lIus"iarus seeds, Iife-lipan of, Z
~ hiflorus seeds, life-span of, '2
- scoparius seeds, life-span of, 3
Daclylis g/omrrara seeds, longevit}" of, 68
Dandelion, sec Tara,ramm QjJiri71al~
nanthonium srmiQ71nularis seeds, longevity of,

'"

Daphne _::.nnnn seeds, life-span of, )

2m

SUD PRLSERVATION A..'ID LONQ.""VIlJ.'

Dalura .seeds, life-span in soil, 9

- - mutation rate affected by age, 15]--6
Daucuf ~arota seeds, effect of relative humidity on viability of, 18
- - - effect of temperature on viability of,

,8

- - longevity in storage 50-52, 54

- - moisture content of, 22
- - safe moisture content for storage of,

"

- - - u s c of old, 14r
Drlphinium ajans seeds, effect of bclowftce'nng temperatures on, 62
- hyb,idu1ll seeds, effect of below-freezing
temperatures on, 62
- seeds, longevity in storage, 55-57, Figs.

8,9
- plants produced from old seeds, Plate .j.
- use of old seeds, 141
Dcschamp.ia. seed" longe,ity of, 68
IXsiCCItion, effect on seed longe\it)', 94---98
Dctcrionltion of seeds
- - - (::luses of, q.&-S7
- - - clIange in alkaloid content during,
-

'5'
-

- change in auxin content during,

152
- - change in chemical composition
during, 148-S0
- - change in inhibiting substances
during, 153
- - change in organic acid content
during, 152
- - change in vitamin content during,

'5 t -z
-

increased mutation r.m: during,

E!cetriC:ll responses, as measure of seed vi:r.biliry, 1)6

Euotharis 'aha seeds, longc\iry of, 92
Eim, Sa Ulmus
l'fymus gll/"'''s seeds, clfect of maturity on
\iability and longc\'ity of, )8
Embryos, development on moist filter-paper,
PI. S

Enzyme acti\;ty, = e n t for tcsting

seed viability, II5-17
- - of deteriorating seeds, 150-1
E,uhtius himujfoHa seeds, life.-sp;ln in soi~

8
E,iotaufon uptanfulau seed:i, longevity of, 92
En!um uns seeds, life-5J"l1l of, 2
E'ythroxylon seeds, effect of desiccation on
\'iability of, 98
J:;uraJyplliS minWla seeds, longe\;ty of, 89
Eurommia ulmoUi~s seeds, 'i:lbilitr test by
uciset! embryo technique, I I )
EuonYT1lus sp. seeds, \;ablli,y test by e<cised
embryo rechnique, II3
Euphorbia ~srola seed.~, life-span in soil, 10
- margi1lilla seeds, viabilitJ test by exeised
embryo technique, 11)
Euphorbiace:le seeds, life-span of, I
European mountain a.~h, su So,bus aucuparill
E);~d embt)'o technique, for It'Sting seed
\-iahiliry, tl2-15

Fagus seeds, terr:a.zolium testing of, 133

- SJ/r;a/~a seeds, selenite test of, 119
Fescue, su Futrlra
Fe>luca rubra seeds, absorption of warcr

,8

153--'7

- - - relanon of enzyme acth'ily to, 150-1

Dicotyledons, vii
Didiuus tatfufta, see T'a&hymm~ Mtfuf~a
Diod~a pauci}fora seeds, life-span of, 2.
Dinirrobenzol, for testing seed ",iabiliry,
11l)-'20
Djpfodia ua~, longe\'ily of, on seeds, 46
Dogstlil, crested, see C)"1I0Sldln crista/us
DoN~hoI IifnoIus, absorption of lllQisrure by
seeds of, 18
- unguicula/us seeds,life-sp=of, 2
Dormancy in so:eW:;, effect on germination
and longe\'iry, 38-39
Dl);ng agents, for seed stonge, 29
- artificial, for seed stoT:tge, :z8-29
- pre-harvest, of seed.., :29
Dyes, for testing seed \;.ability, .lli
E,hillo'Ys/is seeds, ,'i:r.bility test by excised
embryo technique, II) .
Eggplant, su Sofanum mef01lgma
Egyptian mummy seeds, see Barley seeds,
Egyptian mummy
- - - u~ Pe:l, Egyptian mwnmy
- - - sa Wheat, Egypti:m mummy
Eirhhornia rrassipu seeds, germination requirements for, 93
- - - longevity of, 93
Efa#nc amtfirana seeds, longevity of, 92

by~

- - - tetrazolium test of, 124

Fesluca seeds, 10nge\itJ of, 67-68
Field crop seeds, longevit}, of, 63-';0
Sf( alIo names of individual plants
Fiorin, 1U AgrOIlis slownifffa
Fir, Dougi:ls, IU Pseudotsuga tl1rifoiia
Fire e:uinguisha chemicals, effect on seed
\-u.bility, 48
.Flax, su Lillum usitl1/issim/lm
Flower seeds, effect ofbclow-fl"CC'Ling S'Ol":lge
on, 33
- - longe\;t}" in storage, s.;-6z
~fU also individual names ofspccies
Fluorescence me:asurementli, for testing seed
viabilitJ" 120
Formazan, 44, 121, 133
Foxtail, meadow, fU Afopecurus pratenm
Pragaria seeds, longc\'ity of, 7i
F,a:rinus amn-irana ~, viability determined
b}" terr:a.zolium salts and excised embryo
technique compared, 129
- seeds, longevity of, 86-87
- - rule.. for lCSnng with terr:a.zolillrn salts,
135
- viability test by excised cmbl)'o
technique, ll<j.
Fringe-tree, U~ Chionallthus ;,irginira
Fruit, description of types of, \;ii-;:l;
- or pulp effects, on seed longe"ity, 39
- n:prcsentatil'e types of, .Fig, I

202

SUBJECT Ii'il)EX
Fungi, effect on chemical ccmposition of
whe:lt; +I_ effL't't 011 longe,ity of conon seeds, 44
_ cffccr on longtaity of maize seeds, -l4
- effect on longevity of rice sn-ds, 44
_ effect on longe,-it}- of i>I-'Cds, 4G--<{(,
- lifc-l>-pan of, 45--1-6
FlWUium mc~ilifcrmc, effect on dctcriol'lHion
of barley ..ccds, ,p, 1'1. 3
_ - longevity of, on seeds., -1-6

G"uini.. l1Umg~s/lJn .. seeds, effect of dcsicotion on \'i:lbility of, 98

Ga"",,,-,s c-lchange, in old Ntfumb~ fruits, 35
- - in SlOred seeds. 33-34
Gases, dfect on viability of seeds in storage,
33-36
_ see "fw names of different gases
Genetic df=, on viability of OrJZ& sa/iw
stttls, 36
_ _ _ - - Zra mays secds, 36""'37
Ccn/iana mnita sccds, longevill in storage, 61
Gcr.lIliac"llC seed~, lifc-span 0 , I
Geranium Ilissutum seeds, lifc-sp:tn in soil, 10
Germination of see<h, aner sturayc for 150
days at 'l1rious tcmp<:r:ItuTCS and relative
humidities, Fi~. 3
- - - effect of dormancy on, 38
Gibk,ef{a ua" longevity of, on seeds, -1-6
Cflldiutus seeds, !onge,'ity in Stor.Igr, 61
Glossary, 162-5
Glycm" striata l;Ceds, longc\ity of, 92
Gly.inc mu seeds, chan;:-cs in proteins during
stonge of, IS0
_ - -longc'..ity of, 71-72
_ _ _ yields of old ;md young, 139
Compf:rma glabosa seeds, viability lest b}
eu';;;cd embryo tcchniquc, I l J
Goodcni:Iccac sec:d.~, lifc-sp:m 01, 1
Gassypium hirsutum seetls, viability delermincd by IClr:lrolium salu; :md excised
embryo technique ccmpared, IzS
_ seeds, eIrect of age on plant qu"lity,
13<)--40
- - effect of chenllC:I1s on, -t8
_ _ clfCCI of drying on longevity of, 28
_ _ effect of micrj)-()rganisms on, -l--l_ _ effect of sulphuric-acid dclinting on
kmgevity of, 37
_ _ cfTct:1 of ~'aI"iou.~ gases on longc\ity of,
33-3-1- -longevity of, 72/4
_ _ oxidase content of, tl7
_ _ tetraroliuffi tcsting of v:i:lbililr of, 127
Grains, effect of oxrgcn on 10ngevilY of, 35
_ effcct of partial \1lcuum on longe\ity of,
_

3~i:l.bility

re;rcd with tetrazolium sailS,

.21---6
Grass, bluc, su POD pratcmis
6=, French oat, ser Avnlu datior
Grass, Jobn<;on, Set /falem halcpcnsis
Grass, orch:ud. sec Dad)'!is glomcrata
G= secds, longc,ity of, fr]
_ _ ree alsa name;; of individual species
Gymnosperms, \'ii

Ha!,,&clan &!O,"ffalm seeds, longevity of, 93

/l"malllclis tJirgi"i"'la scetls, vi:lbili[)' dctcrmined by tetraro!ium salIS and exeiscd
embryo technique compared, 128
_ - - ~;abili[)' Icst bv ~cised embryo
technique, II3
.
Heating of seeds, effcct on germill:luon, 154-5
- - - elfect on mutation rale, 154-6
- - - resistance to, 154Hdminlhosporium, longe~'ity of, on sc~~s,
-1-5-4 6
Herniae\;:, wcstern. sa Ijuga lutcrophylla
Hcto-anlhrra dubia seeds, longevity of, 92
/leua b,a1iliens;l secds, effect of dcsic~"ti()n
on viability of, 95
- - - tnln~por:ation of, 159
Hibiuus ,,,,,,,abis seeds, longc\i[)' of, 76
_ cuulrotus seed~, wc moisture conlcnt for
storage of, 21
_ syrill'/ls seeds, "iahilit), dctcnnined by
tctr:tzoliurn salIS and excised embryo technique compared, 128
lIofcus ha/~pc",is seeds, Oltalase activity as
measure of viabilit,.. of, 115
H(f1drom t'lltgarc seeds, 48
_ _ _ eytologiClI and genctiol changes in
ageing, 157
_ _ _ elre..:t of fungi on germination and
seedling de\'c1opmcnt of, PI. 3
- - - effect of oxygen on X-irradiated
Sttd$ of, 35
_ _ _ mutation (".Ite affected by age of, 157
- - - selenite rest of, Il9
_ _ _ letnlzoliwn Icst for \;ao,lity of,
Fig, 12, 123, 124, 126, 13-1Humidity. COllStant, method of mainlaining,

'5

HymuwphySIJ pubcurns seeds, life-span in soil,

"

Impcnneabk coats, In Sa:d-coalS, impermeable

Indi:ut 10tllS, su Nclumllo nuc.frra

Indigo carmine for testing secd viability,

117-.8
InhibiUnA" S1lbstancc:s in ageing seeds, 153
Insect infCSt:ltion uf storcd Stttls, 48-49
IpottllNa la,unosa seeds, life---J;pan of, 9
lridcae seeds, life-span of, I
]:raphMUS u"iretus seeds, telrV.olium staining
of, uS
japan-Q!lince, see CharnDmtksjaponica

Juniprrus seeds, nJ1es for tcsting with lemzolium salts, 135

K.aroo bush seed.~, selenite testing of, I 11)

Karhia indica seeds, 10ngcvilY of, 76
_ uopa,ia sced.'<, effect of below-freezing
temperatures on,

6~

Labiatae seeds, life-sp~n of, I, 2

lAala tllti~, absorption of moisture by
s~ds of, 15-:11

23

SEED PRF..SffiVATION A"'D LONGE\'lTY

Lac'urll JlI,ir:u seeds, effect of :I$e of seed on

pbntqu:llity, 138, '41-Z, 14Cl--7
- - - effect of moislure on ,-iabilily of,
!.1.-';l!

- - longevity in storage, 50
- - moisture contenl of, 22
- - safe moisture content for stor.LgC ni,

"-

- use of old, lop

- - - vi:lbility of imbibed seeds of, 13
1,.Jl(f,mtrmiQ "/laif" seeds, mbility tesT by
acised embl10 technique, II]
seeds, Ionge,ity in stonge, 58
1"'[!,fIt1' p~,,"-/lNJ Sds, Iif~ of, :!
Lc:guminos:Ie seeds, ll{e-t;pan of, I, :I, 69-10

u''':p-w

- - sWning of, 126

- )ields ~ old, 139

5(( IZJso Il2mcs of indhidll2l r.bna
upU/ium (CIUd4ria) ibilh.J seeds, life-span in
-

soil,
-

10

rtjau

- JlIti=
of, 120

seeds., life-span in soil,

seeds.

10
Buonsc;:ence ~ 1 S

Lsipinus r.H,guslijofius seeds, :utilici:Ll drying of,

-

'"
-

~"l':S

-!JjrSUlus~,

during storage of, 149

absorption of moisture by,

"

Lyc"pe:nirum =kntum seeds, absorption of

mo~ture by,

15--21
.
- - effect of age of, on plam quality,
41-z, LH--6, PL 12
- - - effect of mois= on \ubility of,
14-2T
- - - clfect of rehti\"l: humidity on \i:I.
bility of, 18, 25-28
- - -effi:a oftempera= 00 \iabiliryof,
-

"

- - - Jonge..ily in s~, 51-5=

- - - moistuTc:.conlcnt of, ZJ, 26
- - - slfe moistun: content for stor.lge of,

"

---use of old, 141

Lyan"tk.rr=-olS .foJriJromJus seeds, 'i.l.bility
tesl by en:iScd embryo technique:, I I~

or,

---:uu.olium treatment
,:I,
- - - viabilitr l=d by use of pon.ssium

bJdroxide,1I7
- ciqjniaun seeds, life-spm of, 7, 8

LespritZ4 mttrmdllJ seeds, life-sp;m in soil, 9

1..emK:e, s WtlK. ~tmr
.l.twtImiJ ~~tPb.21.: sced.~, life-span or, 2
Light, effect on seeds in stOrage, 36
Lif,llSIr..". seeds, 'iability detttmined by
tetnzolium SOllts and uciscd cmbr}"O 1f'Ch-mque compm:d, !:g
LiN"", ..,..,_ seeds, Ion~ity in ~
-

;,
r~lJlr scetb,
PL,

Ionge\iry in storage, 5g--6T,

J..imn.mtUnumt nympAulliJu seeds, Ionge... ily

of,9 2

Lirlum seeds, longe"ity of, U

- :citoTlilsinuUlueeds, absorption of moisrure
by, IS-ZI
-

effect of moisture on vi2bilit) of,

14 21

- -

germinability of, 4 i

Liqlliallmba, Jl;TroTrijilllJ sced.~,

\i:lbility determined by Ictta7.nlium salts and ucised

embryo techniquc compared, !28
- - - ,iabi.lity ttst by excised anbryo
technique, f1]
Liriod-mdrrm tulipi[mJ seeds, effcct nf dl:$ic_
cation on viability of, 95
Liuhi seeds, effccl of desiccalion on viability
of, 98
l-obdia. JOrlmllnnil seds, longevity of, C)2
ufjum peren/le seeds, moislure COIlJellL of. 18
- seeds, lon~"ity of, 68
- - retrarolium testing of, 124
Loniaro dioiciJ sccd.~, ,-iabilit}' determined by
tetnzolium $:fIrs and e:xcised embl)'o tecb.~
nique ~omJXIred, 128
lAJphDfO(arpus spDngiQsuJ seeds. longevity of,

"

Lolus ufitinusus seeds, lif~pln of, 3

Luccmc:, ur MtdicilgO U';r:oT
Lupine, blue, sre l:Jpinus aJl[lJSlijo/iUJ

AfoU<Qwi<l seeds, \-iability of,98

M:t;2, Sre .'If== ,puUI1:issiAl
M.:ow/i. MIalri".ta. seeds, mbility test by
cu:ised ernbt}"o tochoique, IIJ
M:Ihog::my, ~lc:af seeds, dfc:a of desiccarioo on liability of, 9S
.\Lain:, 5(( Zt<l _ys
~bbchitt gr=, for te:sting seed ,iabililY,

,,'

AloTW soed.~, comparison of aQs.ed embryo

md t:etr'.zoIinm lew; of, 132
A[I/m. T"lrDI:Ii[oliIJ seeds, life-sp:m in soil, 8

,\WVaee:lC seeds, lif~ of, I, 2

?'Ungosu:e:n, sa CIIUUU4 1T.4ngIlJliln<l
Maple, sugar, su _-laT mh:zrum
MartplU1. seeds, liability lest by excised
anbl}"D Icchniqoe, ITJ
i\farurity eiT=. 00 the keeping quality of
~3:r-J8
_\ftdiClSgil SIlti;;a seeds, elfecr of scarificalion
on longeriry of, 37
- - - kmge\"ily of, 61)
~ ~ effect of age ofsced Oil plant quality,

'39
,udiJIJtus <ll~ seeds, life-span of, ]
-lutro seeds, life-span of, 2
MlnilduroT 5pp. seeds, \Ubility test O} excised
r:mbl)'o technique, 114

Mercury compounds, elfecl on sds in

storage, 47-.;8
- - for control offungi and inset!! on stored
seeds, .;6-~7
Mesquite, Sll Prosilpis r;dutilloT
Methylene blue for testing seed viaoiliry, 118
Microflnr:l, effect on longe\it)'of seeds, 4G-45
- respiration of, q8
AfjnUl.J glomal/til ~, life-span of,::
i\'Eois= content,. flocruation in,
- - effect on keeping qU:llity of TlUlJXlJCUII1
seeds, ;>.J-25
- - of seeds after s[OI1Ige at \-~riou<; teO>P\'rarures and rdari,"t: humidities, Fig. ::

Oak, sn QU~(IlS
O~"ulJ.~,a bim"i$ sccds, life-span of, 7, 8, 9
OenDll:aa seeds, afftcted by inhibitin;

.Mou.wrc content of Sttds, app:mItus for

making seed moisture coment test, 1'1. 6
- - - - 'Critical' point, 22, 2)
- - - - effect of rdati,"c humidity on,

substances, 153
Oily seeds, longe>ity of, 71-75
Okra, su HilJiuus esculro/us
Old St1s, quality of plants produced from,
138-4i
- - }icld from, 138-40
Onion, su Allium upa
OnopoTdum "c"n/hium seeds, life-span in soil, 9
Opuntia seed~, lon~e"ity of, 76
Oran,,"t': G, for testing seed viability, 118
Orchid, ue Cauleya
Qrs:anic acid contt.'Ilt of uettrionlung seeds,

14-17

_
-

- - - effect of tenlpcr.lture on, q.-I9

- - - effect of y:mety on. 14-17
- - - effect of ,iahility on, 18
- - - effect on yiability, 14----2/
- - - in open stor:Jge, 22---24- - - maximum for safe storage for one
year, :1-:1- - - - S3.fe moisture content for storagc
of >1lrIDUS seeds, :2,
- - - - seasonal effect on, zz, Fig" 4
- - - - Y:Iriety cffect 00, U
- effect" during seed rran.~portation, 158-q
- - on im':lSion of seeds by fungi, 40-43
Mooocotylo:doos, .ii
Monle>;uma spuiosis:;ima seed" effect of desiccation on viability of, 9i-QS
Mould growth, effect on chemical composilion of ,,"'heat scc<ls, 4-1, 4-4- - efftct on viability of seed", 40-46
- - effect on Ietrazolium testing, 126
Mungo (bean), see Phllseolus
MUt:ltion, chromosomal aberrations from aged
wheal seeds, PI. 13
- mie increased by ageing in seed" '53-i
M)"lIsofis SyIVII/i'lI seeds, effect ofbclow-frcczinp: remper.lIUTCS on, 62
j\1y,;ophy/fum seeds, longevity of, 9l
M yr!accae seeds, life-span of, I

Naja> seeds, longevi)" of,

'5'
O,,"xajilponica .""ed~, >iahility detcrmined by
tctt'2zolium salts and excised embryo leehnique compared, 12<)
Oron/j,,111 aqua/icum seeds, longe\ity of, 92
O'y= slIti"a seeds, effect of variety on ,"iahili)" of, 36
- - -longevity of, ')3--<)4- - - pre-harvest ul)ing of, 29
Ory= seeds, Cll'bohydratc etc. changes
during storage of, 14.'), ISO
- - life-span in soil, 9
- -loss in stor:JgG, I59
- - tetrazolimn test of, 124Oxidase activity, in seed~, 15I
- - measurement for lesting seed viabiliry,

"7

Oxygen, effect on longevity of GOSSJ'}ium

91--<):1-

Nandina tiomesriea stl'ds, longevity of, 76

NIIS/",/iuIII naj/u,/i,,,,,-aqtla/uum seeds, longe,it)" of, 9:1Nelllmmum sp~domm, s(c Nelmnbo nurif~a
Nelumbo l"Ifa seeds, life..,o;pan of, 3
- mu:iffTa seeds, impermeable coats of, 4,

rLl
- - -life-span of, 3, 4- - - location of old~ PI. I
- - - longevity of, Q4
- - - vigour of plants grown from 01<1
seeds, I38, PI. I
Ncutr:TI red, for testing seed viability, 118
Niroliana seeds, longevity of>
- - growth from old, 14-
- tabacum sccds, lifc-5pm in soil, 9
- - - mutation rate affeeto:d by :Ige of
seeds, etc", 153. '57
Ni.vospoTa l)1'y::.iU, longe,~)' of, on seeds, 4-6
Nitrogen, effect on longevi(y of GIISSypill/lJ
seeds, 34- effect on !ongel-iD' of 0,)":.:0 IlI/i;;>a seeds,
35
-effccton longevityofua mays seeds, 34-35
.1I,'DlhofIlJUS mrn:;j~$ii seeds, longe,-il}' of, 88
N"pJ.or /"I~ltm ~1'Cl1s, longe,'iry of, 'P
NYII/phaeil alba seeds, 10ngeviD" of, 9:1- luoerosa seeds, longe.ily of, 9z
Nymphac.co:ac seeds, lifC'-Span of, 1,:1Ny",phiJIumlltus varirglllu$ seeds, longe'"ity
of,9 z

is

seeds, 34- effect on longevity of grain seeds, 35

- effect of longc,i~" of Ulmus amernana
seeds, 35
-effecton longevity ofZta mayssa:ds, 34--35
- effect on longevity ofX-irradiared seeds of
Hordeum, 35
Packets, clfecr on seed viability, 9Q-IOQ
- effect on viability of onion seeds, IOI---{)
- type of, for main(aining seed ,iability,

",

Palm seeds, packing and tI"aosport:ltion of,

'58

Panicum seeds, longc"iD" of, 6Q

Pans)", $(( Viola
Parsley, Itt Ptlroidinum iiJli-::um
ParsnIp, su Pas/ina'a satha
Paflhrnorissm quinque/oUIl seo:ds, viability
determined bvtetrazolium salts and excised
embryo teeh';ique compared, 129
Paspolidjum j"bifiorum seeds, longevity of, 69
PII5/ir.aca sa/iva secds, viabiliry in storage, 50
Pea, Eim'ti:ln mummy, viabilil}' of, 5
- garden, SU Pisum sa/....'ltm
- sweet, su Lalhyrus
Pe:!eh, ue PTimus persico, SU a/so Pnmus
Peanut, su Arachis hyjlOgoeo
Pear, sa !'.)'rtlS
Pecan, sa Carp: iflmowsis
Pdlfmtira 7:irg;njca seed~, longevity of, 0):1Pcnicilli""" effect on deterioration of barley
seeds, 42, PI. 3

25

SEED PRESERVATION AND LONGE,'ITY

Pepper seeds, safe moisfUT<' contall fo r

S~of,21
- - IC( 41sD C.psiautl [nUlCtrrl

Pcroridasl! in stals, 151

P("olClirrum lIIrit-u". seeds, effect of bclaor.
freezing temperature on, 33

PmtnUl. nyJJrid4 seeds, effect of bclo,..~ft=T.illg

temper:l.\ute on, 33, 6z
PIUlUOfuI seeds, anifici:l1 dr)'illS of, 28
- - eflect of age of seed Oil plallr quality,

13~.P,

PI.

10

- dfeet of rebti'-e humidity on ,iability

of,18
- - effect of tcrnpmlrurc on viabilirJ of, IS
- - Iongc:vity in stonge, sJ
- - safe moisture content for stonge of, a 1
Phenolase aahiry in ~ 151
PIdautt prtllOUt seeds, Ionge\iry of, 68
- - - testing \iabiliry of, 136
PhJUDlItlcltp seeds, effect of dcsiec2t:ion on
,-iabilit}' of, 9S
PhyfoUuftl ilmaXana seeds, life-sp:rn in soil,
9
Piaa "biu!lt.ocds, effect ofmaturit)'on keeping
quality of, 3S
- - - test for viability of, 117, 119
- HmaJfmil, IU Piua Zlali(a
- CICCW seeds, effect of below-freezing
storage on, 3'
- ZUlliftl seeds, e!fect of bclo""---fu,::ziug:
storage on, 31
- seeds, comparison of germination, excised
cmbr)1l, ~ tcmzolium tests, 130
- -Ionge",q' of, ,S, f9, 80, 81
- - \-ubility ~ by use of indigo-cu"mine, 1I7
Piua liJcnrrrm seeds, effect of below-freezing
storage on, 31-33, Fig. 6
- - - eff~t of moisture reduction On ,;ability of, 33
rine, loblolly, let Pinul/ada
- longleaf, IU Pinus pafuJ/ris
- pitch, Sa Pirrlll rit;iJu

- ponderosa,

ICC

PitJus pqndrrollJ

- red, ICt Pi",11 rcmr"Sil

- Scoa, ICC Pilllll l]k61ris
- shortJeaf, ICt Pi,,," amlltlla
- slash, ICc Pirrlll CtlribUil
- sugar, ICC Pinm 4utrbo1Ul1l4
Pmus critl(tI seeds, dfca of beIo'W-fnx:zing
SlOf1lge on, 31
- - - eff~l of moisture and tcmpmlture
-

on,~
cembra seeds, rules for testing 'With tetn-

zolium sailS, .35

uhinilltl seeds, df~'tt of bclow-free7illg
storag<: on, 31
-lanlbrrlUl.tlll, cone and winged sec<.l uf, Fig.
-

lIiKrll, grDl"th of ncised embryos, 114

- selenite lest: of, 119
- pslllllriJ seeds, absorption of moisture b~..
T5~

3'

- -

- effect of bclo..---freezing stonge on,

effect

~anent,

110

of 1o'W-lcmpcnrure

pre-

"urutris Sttds, effttt of lI'lOisrure on

.iabilitr of, 1-4-21
- --loss of,iabtlity by, &>
- - - mcltod of resting \uhility of, I I I
- - - moisture amlClll of, 2~
- potJ:krOJJl seeds, clfect of below-freezing
storage on, 31-33
- rcsinosa seeds, effect of belo...rn:cUng
storage on, ]1
- fjgida seeds, de"c!opmcnt of embryos on
moist filter-p:lper, PI. 8
- seeds. catalase acti';t)' as measure of
,-i.ability of, 116
- - oompuison of germination, ucisro
embryo, and tetrazol.ium rCSlS., 130-1
- -Iongc>;t} of, 7S, 79, So, 8., Rz, 8]
- - viability teSt by ac:isc:d cmbr)"O
tcdmiquc, II~
- - nabilit) 1C$led by use of indigo carmine, 117
- 11/~ seeds, ripening of, :;8
- - - tQt for \-ubilit) of, 117
- - - X-ray analysis as measUTC of ,-iahility of, 137, PI. 9
Pinus rada seeds, effect of bdow-fn:e1Jng
etc. stonge on, 31, 80---82
- - seedling produ~1ion from s-ycar-old
seeds, PI. S
Pinnn ltIlirum, pods and seWs of, Fig. 1
- - seeds, safe moisnrre contCilt for storage
of, ::l.I
- seeds, Clt2bse 2Cti\"iry :as me:zsun:, of
\iability of, 116
- - mnt2tXm nrc affected by age of seeds,
1 57
P1JUI1alllllllJp,r seeds, life-span of. 7, S
Pbsmolysis, for testing sted ,-iability, 115
Plum, sc~ PrUtlUI
PIN' pratnlliJ seeds, longevity of, 68
Polygonaceac seeds, life-span of, I
Pol,Kanum h,Jfo,ip(r seeds, life-span of, 8
- seeds, longevity of, 9a, 93
Polyploidy, .n......S
Ponlcderia C01'dllla Sds,longnity of, 92
Poplar, ICC Popullll
Pllpulus seeds, effect of :Ur on Iongevily of,
Pi1l11S

3l

- - effect of oXJ!m on 'liability of, 95

- - Ionge,ity of, 87-88
PffflW=-iJ 1Ikr'IIUIJ seeds, life-span of, 7, S
Poz/U1log~ton seeds, Iongcnty of, 91-i)Z
Porassium hJdroxide, for testing sted ,iability, 117
Potato, It: Solanum tuh(f'orum
Poren/iffll nort"tzica seeds, life-span in soilg
I>rimula mlll/lcuMu seeds, dfe~t of belowfreezing temperatures on, 6a
PrinlCpia sinrrrsil seeds, comparison of e:t:ciscd cmbl)1land tetrazolium tests of, 'J2
Proltlpis luplumitlrr" seeds, emf tre:ltment of,

,.

- ~bnituJ seeds, Iongenty of, 76

PnmI/.J i:merietl1I4 seeds, do:lopmenl of
cmbnos on moist fiher-p:>per, PI. S
~,ity of, 88
_ persUil seeds, effect of pulp disintegration
on ,iabilily of, 39

- --=-

206

SUBJF.Cf ll\'DEX
Rosa muftiflora seeds, comparison of excised
embryo and tetrn7.olium tests of, 132
- seeds, rules for testing with tetrazolium
5:llts, 135
- - viability tcst by excised cmbIIo technique, TI.j.
Rosaceae seeds, romparison ufexcised embrJo
and tetrazolium ICSts of, IJ2
- - sto",ge of, 88
Rubber tree, see Hroea brasilinlii~
Rudheckia kina seeds, life-<;pan ill soil, 9
Rum~x ,,/spus seeds, life-span of, 7, 8
Rre, see Seca!e ~er~afe
Ryegr.JSS, sa LofiufIl

Prumu seeds, comp:ui;on of c:xcised embryo

and terrazolium {ests of, 13:&
- - rules for testing ,,"ith tctr=1ium salts,
135
_ viability tests by excised emhryo
technique, [14
hudolSuga t",riftlia seeds, comparison of
gennination. excised emhrJo, and tCtnlwliuffi tests of, '3f
_ - - development of embryos on moist
filter-paper. PI. 8
- - - effect of below-freezing storage on,
31 -33
- - -longevity in packets, 107-9
- - - viability test by excised crnbf}-rl
technique, ll-l- - - ,iability tested by use of indigo

Saunarum seeds, e!feet of desiccation on

,-iabili!)' of, 95
Sagittaria seeds, longevity of, 91, 92
Salix seeds, longe"i!)' of, &7
Salvia splend~nsseeds, effect of below-freezing
temperalUTCS on, 6z
Sa/urtja hflrtensis seeds, viabili!)' in storage,
50
Sdrpus seeds, longcvity of, 91, 92
&vt~ pine, su Pinus syl'Vt5/ris
Scaling, method of, cfftet on keeping quality
ofTllra:rllnim seeds, "3-25
Scra/e rena/e seeds, mUbtion rate affected by
age of seeds, 157
_ _ _ tctrawlium test for viability of,

carmilH:, 117
Pul~a isopbyfla seeds, viability determined by
tt:tTazoliuJn salts and excised embryo tcchnique compared, 129
Pumpkin.,

jU

CI'-'Iffbira ptpr

Purrhia tridmtaM seeds, viability (cst by

e:xcio;ed embryo technique, 115
P)'Tm seeds, comp;ui<:oo of excised cmbrro
and terramlium tests <;Jf, 132
- pomc and seed of pear, Fig. I
- seeds, \iability test by e:o;ciscd embryo
Ilxhnique, 1'4
~eTf"S seeds, ch:mgcs during stolOlge of, 149
- - effect of desiccation on viability of, 95
- - tetrazolium testing of, 133
Q:!dnce, 5te CydQllia Qb/Qnga
OJllirine, sa CinchQna kdceri4na

Fig.

Radiocarbon lhting, to determine age of

seeds, :;
Radish, see Rapf.alllls sath'lIs
Range grJSS seeds, dfct.1 of maturity on
viability and longevi!)' of, 38
_ - - sa alsQ scientific Il:l.II1CS of grasses
Rlmunrolus seeds, longevity of, 92
Rapkallli5 $l<livus seeds, fluorescence m=remcnts of, 120
- - -longevity in storage, 53-54
- - - resi.tance to heating ol~ 154
Records of old sccds, 1-6
Resazurin, for testing seed viability, 120
RhamnuJ jafJIJnirto seeds, viability determined
b, temwlium salts and excised embryo
technique compared, 1~,9
RhQdo/ypos lWrWidu seeds, dC\'clopmem of
embtyos on moi.t filter-paper, PI. 8
- - - viabilicy test by excised emhr}'o
technique, "4
Rhus arGmlJlica seeds, viability determined by
Il:trnzaljum salts and e.",cised embryo tC'l;hnique compared, u8
Rice, U& Or)';;4 Silli;;a
Ricin", rt1mmrmiJ seo:<.b;, lifC,.;5pan of, 6
- seeds, longe,icy of,
RGhini:l puudDlU:lcia seeds, life-span in soil, 9
- - - ,.",bility determined by tetrazolium
,."lts and excised embryo leclmique compared, Iz8

'is

12, 124

Sccd...coats, impermeable, cffccr OIl germin:ltion and life-span of scccls, 37-38

- - - of Legnminos:lc, 4
- - - of Ndumbo nli~iftra, +. IJ
Seed industry, ix-x
Seeds, deyelopmcllt of, vii-viii
- buried in soil, life-span of, 7-13
- macrobiotic, 1
- mesobiotic, I
- microbiotic, 1
_ representative, Fig. 1
- of culti""ted plants, ,-iability in soil, 9
Selenium salts, for testing seed "lability,
118-1 9
Scploria nodomm seeds, longnity of, on seeds,

45
Sequoia (Sequoiadendron) gigantca seeds,
longevity of, 79
Shcpherdia argmt&iJ seeds, ,'iability test by
excised embryo technique, 113
SUme nlJ&/ijlorll seeds, life-span of, 7, 8
Sinapis alha seeds, tluoreseence measurc"
ments of, 120
Size of seed, as rdatcd to life-span, 3
Snapdragon, SC& Anlirrhinum majm
SQla"um taro/menu seeds, life-span in soil, '0
- daeagnijofium seeds, life-spm in soil, TO
- mtu}ngcnto seeds, 25
- - - longevity in storage, 50-52
- - - moisturc content of, 22
_ _ _ moistl1fC content of, at yanous
rcb.tiyC hwnidities, 26
---useofold,I{1
- - - \il.bilitv of, at various relati,c humidities, ::5--28"

20']

SEID PRESERVATION A.1\:D LONGEVITY

Sulllllilm lIi!rJim seet.I;,;, lif<:-span in soil, 9

~ {IlDa-owm seeds, longc\ity in stonge, 54
Strrtn.s aucuparw seeds, compuison of excised embrJ'O .nd tetruolium teslS of, IJ?
~ - de.elopment of embr)os on n""",!
liIter-paper. PI. 8
~ - 'iabiliry tCSt hv e~ rntbf)"tl
teclmiqne, I r.J

Strrzlrll5lnml seeUs, IoD~\;ry of, 69

Suuthem.Bc:tthes, 1 NOfMfips [mm.usii,
etc.]
So)':l ban (Soyb=n), ICC C/ymu m.u
S/fjrzlllli"1fI nufKartum seeds, vUbiliry rest
by excised em.b~o technique, 114
- seeds, Iongenry of, 9Z
Spap/...rtY1Inl seeds, Iongcvity of, 68
Spe:rnutophflCS, \ii
Spinach, 1 Spiluzriz a/cro:cu
S,mNU ala-au. ~ effect of rmm'C
humidity <WI \"'bi1i~
18
- - - effect of tempentufe on \iability of,

or,

,8

- - $lfe moisture content for S!on~ of,

"

Spruce, Norny.

In Piull

liM

- Silb, sa ~(l1lit(lwrosil
Spun).. ICC Spapu ruroWs
SqUtib, buucmm, In C"fltt"W11I
SI~"yS 1Uldif9lill ~ life-spoul or, 2.
Staining methods for tt:liting seed .u.bility,
no

Stcralliacc:ae seeds, life-span or, 1

~uL1. seals.. elfeer of nutwity 00
,-lability and Jongevit]' of. J8
SriZilWi".. s.eeds, Ionge\il}' of, 70
Srongc c:onditioos, determiII1tion of Iong-

Slip

evit) by, 157

f;IQlities; in ''lIIriOIl$ OOIIntrics, 159'""'61

-llJliKS,. 160
- of seeds, necessity for. '"-'\;
Stratificali<ln, method for Iow-temperarure

pmreanncnt of seeds, 95

Strawberry, ICC FrllZaria

SUIfll" beet, Kt Btf.. t"flf!aril
- --ane, 1 SiUrMnim
Sulplu drugs for contrOl of fungi in stored

seeds, 48
Symphorirarpos

ram,,~sU1

seed$, "i:lbili~'

test by excised embryo technique, IIJ

TaraxllC1lm o.fficinal~ $eeds. effect of moiscure

conTent on keeping quality of, 24-25
- effect of moi~ture Jlucru~tion on
keeping quality of, 2;;--2.'i. 28

'I'tn;odium distir!luffl seeds, longe\'ity of, 79

TllXus seeds, rules; for testing with tctr~zolium
salts, 135
-

- ';abilit)' determined by teO'llzolium

salts and excised embryo techniqne comp"red, 12<)
Tellurium salts, for testing seed \'iabilitl' 118
Tempcr:1cure. below.frcC1ing for Ill2mten_
:mce of \"i:tbility. JQ-J3
- during set.-d tr.I.nsportalion, 158-9
- effect on longc\i~. 30-33
- relation to moilltur<: :rnd longe;ity, 30

Tetrazolium salts, for tesling s.::.,d ,iabilil,.

J21--J6
- - for lo:stiD.g 5Cel1 ,-iability ~'Offipa.rtd wilh
e:::tcised embryo technique, t2S-JJ
- - mecluorsm of :IIction for testing seed
liability, 133-1
- - rules for usc for testing \iabilit, of
scctk, 135-6
Thiamin in storW wbc:a! gnin, 15f--2
Thuj.. iKddmJaJis seeds, compari~ of
~ ~ embryo, and tetr.lzolium tests, IJI
- p!ic4ta seeds, e!fecl of bclo....frcezing
S!'Onf!e on. JI-3J
Th}IDCbe2cae seeds, lif~ of, J
.
Tilill seals, rules fOl" lesting ..'ilh Ictrazolium
nils, 135
~ seeds, life-span of, 1
Tunoth~-, '" P!IJcJl. ~i"t>IK
Tob:zcco.:Irt "';/&QtUuu
Tomato, sa L~'lml tsntlC%t:U1t
Tone sulNanas in a~ 5C'tds, lil
Tr.uJrp:cu cturuk~ seeds., nabilitJ lest by
arised embryo rechnique. I q
Transportation of.--<ls, oonditions foc, 158--v
- - - imjlOlUnCC of moistun::IInd temperalUre in., 158
Tra.1M BJtions seeds, Ionge,il}" of, 92
T ~tme:m of ~ cbemicals for contrOl
of fungi and imccts, ..;6-";9
Tree seak, longe_it}" of, 78-i)o
- - >Ubility u:sted with teuuolium salts,
127-33
- - .-iabili~- Iesl.ed by X-l'lIy analysis, IJ7
Ut ab4 IlaIJIeI; of indhiduaI 'pecic:s
Trifofjrnn ~u $eeds., lik--$pa.n of. ::
Uu:1lTIUU1ml seeds, artificial drying of, 2.ll
j!T1J1t=e seeds, effect of age of seed on
plan! qualilJ, 139
- - - life-span in soil, 9
- - - life-span of, 3
- upms Sttds; life-sp:m of, 8
- seeds, Iooge"ilJ of, 69-70
- I1ril1trm: seeds, life-s~n of, J
T ri!iom: sati= seeds, effect of mierofICIr.I on
longe,it,.- of, 40-
- - -longe\ity of, 63--65
- - - mUt:ltIDD I":I.te affected by a:rc of. 157
- seeds., ch~nges in proteins etc., during
storage of, 1';0,151
- - chromosolItll aberr:J.tiOIl$ from aged,
PI. IJ
- ~ cytological and genctial changes in
3geing, 157
- - cffecl of3ge on p13nt quality, 139, 148,

'49

radiosensitivity increasing with age,

'53

tetrnzolium t~"1 for "iabiliry of, Fig.

126
- flUlgart seeds, catalase activity as rTH:'2Sure
of ,':iability of. n6
TsrWJ ranadrnsil seeds. comparison of ltttmin:ttion. uci5ed emoryo, and tetr.tlolium
tests of, 1]1
- - - vi:llbility test by e.Ici$cd embryo
tcc:hnique, Tl;j.
-

208

12, 12.J, I2";,

.,,-. :. ' .

.
.. :::.
- ...
'.

SUBJECT lNOH
Viability of seeds, statistinl oompari.>On 01
re,<;ults betwecn excised embryo and tetr:lzalium tests, 133
_ of vegetable ~~~, effect of moisture
oontcnt on, 5:z...-53
- - - - elrect of temperature on, 52-53
- - - - mctbods of testing, 5 I
Vici<> villo$u seeds, viability of 'hard', 31
VIp.,: rintnris, destruction of ascorbic acid
during storage of, 1;2
Viola seed.~, longevity in storage, 58
- tri.o!or \'"Jr. maxima seeds, effect of belllwfreezing temperature on, 33, 62
Virlmin content, of detcrionting seeds, 151-

Tsuga hturophylfa seeds, clfcc[ of belowfreel'.illg stor;,ge on, 31-33

_ _ _ eflect of moisture reduction on
viability of, 33
- - - longevity in packets, 107--<}
Turnip, su Brastic!>
TUISjlago farlar" seeds,longevity of, 16
Typllll [aEiJo/ia seeds, longe,it)' of, 92

UTmus seeds, effect of storage all ,iability of,

Fig.

10

-lan~yity

of, 83-86

- viability test by excised embryo

technique, II~
Umbclliferae seeds, "iabiliry tested by use of

Water hyacinch, S,t Ei.hhorniu ,ri$sip~s

W;l.tenne!on, sec Cilrullus vulgaris
Weed seeds, Yiability in soil, 7-13
Wheat, Egyptian mommy, viability of, oj.
Willow, sa Salix
Wydhiu S&ilbru seeds, ,iability tesr Ill' excised
embryo technique, 113

indigo C<lrminc, 117

Vouh,ium seeds, longevit)' of, 71

Vacuum, parti.al, effect on longe,it)' of Asta

seeds, 35

- effect on longevity of Conifer seeus,

35
- - effect on longe\iry of gr:lin seeds,
35
- _ clfcct on long:c~ity of Ory:;,;tl Sillivo

X-ray analysis, as mca~ure of seed viability,

136-7

seeds, 35
_ _ effcct on longevity of Populus seeds,
35
_ _ effect on longevity of [flmuJ amtrjranll
seeds, 35

Ycasl, fermenting, st.lincd bytetr.lZolium, 121

Zannirhtllia palu!.Iris seeds, longevity of, 92

ka mays, effect of age of sced 011 plant

_ _ clfcct on longe"jty of Vubma seeds,

35
Vallirnrna american,. seeds, longevity of, 92
Variety effects, SU Genetic effects
Vegetable seeds, effect of bdow-frttzing
stOnge on, 33

quality, 139

- - sews, m-tificial df)ing of, 28

_ _ _ cat.ll.ase activity as measure of VL"lbility of, 116
_ _ _ effect of relative hnmiditv on ,iability of, 18
.
- - - effect of storage on ~ugar:s in, 149
- - - effect of temperature on viabili!)' of,

-longHity of, 50--54

j U aho indi\idu:J1 mmes of species
Velvet-be:ms, u, St;::;olobium
Vmidium seeds, longe\it}' in stonge, ;8
VtrhauUI1l hlattaria seeds, life-span of, 7, 8
- thapms seeds, life-5pan of, 8, 9
Vtrbmo seeds, effect of oxygen and partial

"

- - - clfect of variet)" on viability of, 36--37

- - - effect of \'anous gases on longe\iry
of,34-35
- - - electrical testing of, 136
- - - genetic effects on longevity of, 66
- - - grain of, Fig. I
- - -longe\icy of, 6S--{ry
- - - protein changes during storage of,
150
~ - - respiration of, 148
_ _ _ safe moisture conlent for storage of,

vacuum on viabilirv of, 35

age of seed on plant
quality, 141, '43
_ vroow. seeds, effect of below-freezing
tempera.tures on, 62
Vemal-gnss, sweet, f.U Antko:r:anlhum oJor-

_ twcrioidu, effect

of

utum
Viability of buried seeds, 7-13
_ of seeds, effect of moisture content on,
14,-21

"

_ _ _ effect of variety on, 1--6

_ _ -exd~ed embryo technique for testing,
-

IIz-'5
-

methods of testing, 110-37

_ _ _ tc=olium test for viability of,

tzt--2, Fig. 13, Iz6, 134
Zinnia tJrgans seeds, effect of below-frcc:zing
tcmpcntures on, 62
Ziunia Qlfuatjell seeds, longcvity of, 91 93

29

:'"~ :::"-:
:~~ "':'
" "'-

_... . -

.
..
. , '., -

:1; "~~~-

.,

." "

.~ UTHOR

ibbou, ~. V., 96

" "

INDEX
Benne, F.. J., 174
llennetc, N., 125

Aberg, 1::., 5
.'\etoll, E. H., 150
Adams, J. D., 35
Afanasicv, M., 76
Ahlgren, G. H' I 37
Akamine, E. K., 7~, 89, 160
Aki, S., 54
1\lbo, G., '5[
Alcock, A. \V., .j.O, 67
Allen, G. S., 33
AI15Chul, A. M., 48, '7Z
Amato, F. 0', 157
Andersen, A. L., '74
Anderson, J. A., 40, 67
Andren, F" ';7
Anthony, K. R. M., 73

Bemal, D. M.,

12.j.

Bethmann, W., 119

llhati:l, D. S., 150
Bibby, K. M., 88
J3inkley, A. M., 1(10
Dlack, M. A., 47
Blakeslee, A. F., 15;;. 171
Blum, P. I-L, 67
l:llumer, J. c., 78
Hlyn, A. G., 3
Bonnier, G., 6
Booth, R. G., 197
Bonhwiek, H. A., xi, IQ.JBoswell, V. R., IH, 53, 65
llr;ldley, A., 63, 64
Bnnd, C. J, 160
Brase, K. D., HH

ArJ.i, M., '}

Armolik, N., 42, 43
Arndt, C. H., 74
AsakalV:l., S., 133
Atano.soff, D., I5-j.
AtkiM, W. R. G., 18
Ayery, A. G., 153

Brett, C G., 47
Brewer, H. E., zS, iO
llriwn, F. R., 53, 8g
Brocq-Rousscu-Sl"e Rousseu, N. BrocqBrodsm,

Bailey, C. H., 64
Bailey, S. W., 49

lliird, P. D., 126

Baker, D., '32
Bakke, A. L., 67
&ldauski, F. A" 17,}
Baldwin, H. I., 79, 8], I lot

lJaIl, G. J., 58
Balu, V" 161
Barner, H., 8]
Barrett, M.. S., IIZ, 113
Barrons, K. c., 54
Bartlett, L., 138
Barton, L. V., vi, II, 14., 18, 21, 25. 28, 30,
Jr, ]2, 3], 35, 36, 50, 51, 55. s6.. 58, 59, 6r,
62, 79, So, 82, 8+, 85. 87, 1m, go, 93, 96, 97,
100,101,10,,",106, IOj, IIO, Ill, II2, II.J,
141, 14-2, 171
Barron-\Vright-ue Wright, E. C. BartonBasak, M., '76
BaL~ Co G., 79
Battle, W. R., 37
Bayfield, E. G., 152, 172
llcal, W. ]., 7, 8
Beanie, J. H., 50, 53, 74, 75, 138
Il:querel, P., 2, 35
Beinar, W., 46, 75, 1.1.0
llelajc'"", G. M., 186
Ikll, \V., 196
Hen;, G., 124

8.,

II 5

Brown, E., 9, 14., 46, 100, lor

Brown, K 0., IO
Brown, F. R., 53
Brown,]. c., 126
Brown, R., 3
Briicher, H., l20
Brunson, A. M., 66
Buglia, 149
Bulat, H . 124, 13.5
Bunker, H.)., 5
Burgerslein, A., (Ii
Burgess, J. f~., 71
BUI"lison, W. L., 71
Burr, H. S., 136
Bussard, L., 67
Busse, I., 88, 95
Burr, J. L., 28, 70

Campbell,]. G., 152Candolle, A. dc, I

C.a.rnnhcrs, \V., 50, 63, fry

Caner, D. G., 160

Caner, E. P., 6.:;:
Carl1edge,]. L., '53, 154, 155, 156
Cam:er, J. L., 16r
Champion, H. G., 79

Gilds, J. F. 1.., 97

ChiI1<o\"skii, V. 1.,75
Christensen, C. M., x, 40, 41, 43, li9, ,S5,
187, 192
Chriuidis, n. G., 28, 7J, 139
2'0

..'

"

"

"

AlTl'lIOR 1l\'Dt.\:

C..hu, A. C, 98, 152Ciesbr, 1\.,78

CiIf~Ti, R., 5
Orkovskij, V.I., l.j.O
Oarl, B. Eo, 148
Clark, C. F., 54
Dayton, E. E., 46
Demeot, E. D., 59, 61
Cobey, W. W., 75

Cohen, A., 188

Cohen, Hcl1cr- -"Sri I klier-Cohen, O.
Coile, T. S., 79
Coker, W. C, 78
C..ole, E. W., 44Coleman, C. H., H
Coleman, D. A., IS, 152
Coleman, F. 8., 106
Colon, Rodrigue:l- -u~ Rodriguez-Colon, F.
Condon, ~.,t. Z., 98, IOZ, 166. 183
Corc)', V. L., Ill<}
Corron, R. T., 47, 49
Camdl, H. ]., 123
Coulos, C. J., 6g
Cowgill, W. H., 90
(mig, W.!N., 59.61
Crane, I I. L., <)8
Crnwford, J. H., 5+
Crocioni, A., 139
Crocker, W., ~i, 10,38,88,1', l12, 115,
149, lSI, 'S{
Cropsey, M. G., 161
('..rosier, W. F., 47
Crossley, D. 1., 83
euender, L. S., '79. 192
Currin, R. I~., 75. 138
Curtis, J. D., 83
Cuder, G. H. t 37
D:11sl.:0'l', F., 83
D'Amato-su _"-maw, F. D'
Dameron, W. H., 1S9

Dandr. J. E., 3

D:1Ilingron, H. T., 7, 8
Darragh, W. II., l)6
Darrow, G. M.; 77
D:usie, M. L., II.5
Davies, P. W. A., 124
Davis, W. C, 1.51
Davis, W. E., liS, 116
de Candolle-uc Candolle, A. de
Decker, A. E., 74Deeter, W. T., 57
Degen, A. von, 159
Dclg:ldo, R. F., l)S
Iknr, T. Y., 83
De-:ter, S. T.,.53
lJiel:snn, A. D., x, 4Z
Dickson, J. G., 4-3
Dillman, A. C., 74Dimitricwicz, N., If7
Divine, J. P., 180
nixon, H. H., 154
Oomenigg, Sorgcr--uc Sorger-Domerngg,
H.
Dore, J., 94Doren, c.. A. van, IiI

Doroshenl:o, A. V., ".5, "7

Oorph-Pctcrscn-uc Pcrcrwn, K. DorphDoty, D. M., 149
Oo(7.en\;;o, A., 166
Dnkc, V. C, 138
Duchon, F., 11.5
Duffee, F. W., 66
Dungan, G. H., 4-6, 66, 139
Durrell, M., 189
Dutcher, R. A., 184QUit, B. K., '59
Duvcl, J. W. T., 6, g, 66, 93
Dymond, J. R., 100

Earle, T. T., 178

Eb~, Schjcldcrup- - f t t Schjclderup--Ebbe,

T.
Ehes, K., 119
Ebihara, 1'., 182
l':dmundson, W. c., H
Eggebrecht, H., Ill)
Egglc:noo, H., 161
Egorova, A. A.., 177
Eidmann, F. E., 118
Eliason, E. J., 82, 107
Elliott, c., 173
El-Shishiny-see Shishiny, E. D. IT. ,1Engstrom, A., 87
Emfeev, N. G., 151
Esho, H., 68
Esdom, I., 76
Evans, H., 98
bTJrd, R., 133, 137
EI'l"einova, T. N., 151
Ewart, A. J., I
E:<ell, A. W., 3, 4Pabrini, I'ohloudcl.: -$ee I'ohlolldek-Fabrini,
R.
Favilli, R., 123
Fcllol'l"S, H. c., 18
Fcmhol7., D. F., 183
Ficl:, G. L., 136
Fifield, C. C, 150
Filutowicr., A., 4-6, 75, 14-0
Fink, H., 124, 134Fi.~her, D. F., 170
Fiske, J. G., 166
Fleischer, F., 88
Fleminn, F., 112, 113, II+. 127
Flores, F. B., 73
Foy, N. R., 67
FrJrnptoo. V.L. 182
Fnol:, W. ]., vi
Franlcnfe1<l, J. C, 49
Fraser, M. T., 136
Freudl, A., 38
Freyer, E., 1.52
Gabrielir-Gclmood, Ch., 119
Gadd, 1., 118, IIQ, IH
Gain, E., 151, 155
Gane, R., 18,68
Garber, R. J., 4-7

2"


LONGEVITY
Gardner, F. E., 39
G:m!ner, R., 189
Gardner, R. C. n., 39, 95
G:mn:m, II. R., Lp
G:!rncr, F. Il, 53
Geddes, \v. r., 72, 1:20, J.j.9, 152, IJI, 179,
185, 11l7, 192
Gelmond, G:!brielit_ -see Gabrielir-Gel_
maud, Ch.
George, E. )., 83
Gerassimova, H., 186
Gerdes, F. L., 74
Germ, II., 72, IJ5
Gcrsdorff. c..c. 1"., 150, 180
Ghosh, '1'., 47
Giersbach, J., 6" 88
Gilman, J. c., 40, 190
Gisquct, P., 157
Giihin, A., !l3
Gorn::ilez-Rios, 1'.,98
Goodsell, S. F., 45, 125
Gordin, A., 188
Gorman,.E. A., IQ.+
Gonnan, L. W., 69
Goss, \\'. L., 9, 54. 57,58, 100, TOI, lID, 170
GI:lcanin, M., 117
Gr~nstriim, G., 191
Greenwood, R. 1\1., (>9
Gries, G.1.., 139
Griffiths, A. E., 53
Grimm, 1\1. G., II7
Grisch, A..., It7
Gro,'cs, J. F., l.i9, 154
Guillallmin, A., 35
Guillcmet, R. 75
GlInrhardt, R, ;<i, I53, 157, 166
Guppy, H. Bo, 92
Gul"t'witseh, A., II9
Gl.1sLa[wn, P R., 120
Gusta[sson,
157, 185, 191
Guthrie, J. 0., 179. 188

A.,

Haack. 0., 78
Halx:r, E. 5., 66
Hacl.:1eman, J.
171
IIafcrkarnp, l\1.., 35, 177
Haferkamp, M. E., 67
Haigh, J. c., 54, 93
I-Ialfon, A., J20
Hamid, M. A., 37
lbo, K.-S., Iii, 118
H:u:rington, G. T., 112, II5, 151, 154
Harris, R. H., 65
Harr!wn, B'i" 53, 157
IJarru;on G. ., 73
Harrold, M. E., 58
l-laut, I. c., 39
Heinrich, 1\1, 18
Heir, C. E., 58, 82, 107, 1t3, 1J4
I-Icl1cr-Cohcn, 0., 120, ISS
Henrici, M., II9
Hibbard, R. P., 120, 136
Hilberr, G. E., 126, 149
Hildreth, R. J., 160
Hilf, R., '33
Hill, D. D., fig

c.,

Hiltncr, L, 120
Hitchcoei, .'\. E., 93
lIiIC,.a. c., 1i2
Hitier, H., 176
Hoefle, O. 1\1., 185
Hoffpauir, C. L., 152
Hogan, J. T., I8{
Holman, L. E., 160
Holmes, A. D., 39
Hoover, M. W' J '52
Hopkin.~, E. F., TOO
Hotchl..i;;.~, J. E., 93
Huwe, R. W.,49
Hrneiar, G., 97
Huelsen, W. A., 65
Hue)', G., 177
Hughes, H. D., 119
Hummel, B. C. W., {O, 44
Huss, E., 83
Hyde, E. O. C, 65, 12-f
Hyde, M. E., 49, 65

Is:I:lc, L. A., 31, 79. 8z

bell', D., 91, 125
lssatchenko, B., 117
Isshiki, S., TSI

IZ:lrd,

c., 176

JaniSeyskii, D. E., 87

JCII5CIl, C. 36

Jensen, c. 0., '33, 134, 152, IS{

Jercel, D., IZI
Johnson, A. G., 154
Johnson, J., i5
Johnson, J. A., liZ
Johnson, L. P. V., 85, SS, lIS
Jones, D. B., 150
Joncs, D. L., 76
Jones, I I. A., 94, 95, 96
Jones, J. W., 36
Joseph, H. C, 50, 87
Jozcfowicz, M., 154-, 155
Jucl, I., 152
Justice, O. L., 70, 93
Kalashin\:ova, .M. I., 19Y
lSarper, R. F.., i6
Kashar:J, Y., 53, s:;
Kataoka, T., 9
Kal;t:, R., 185
Kearns, V., 6S
Kevorkian, A. G., 81)
Khudairi, A. K., 76
Kidd, F., 95
Kiesselbach, T. A., 28, 139
K.il.:uchi, M., 148
Kincaid, R. R., i5
King, G. 5., {8, 172, 183
K.ir~p;tl:Tiek, II., Jr., 178
Kjacr, A., roo 83, lIO, "9
Knecht, H., r r7
Knud.<;on, L., 61
Kobler, R., IT7

212

..- -... .'... ..:.':,-.

.~

.,

..

-~

AUTIIOR' il'\'DEX

Koehlct, B., 46, 47, 139

Kondo, Moo, 6, 35, 53, 83,
IST,15 2
Kopei1:ovskii, V. Moo, 74
Kornfeld, A., II8
Kostromina, A. 5., IllI
Koyama, 1.'1., 83
Kreidow, K. \V., 47
Kreto\'ich, V. L., 67, 149
K:yer, Too, 8g
Kroeger, H., I8g
Knititeh, Moo, 114
Krueger, W.
19'
Kugler, I., 120
Kuhn, R.., In
Kullco, B., 151
Kundu, B. C., 176
KIlll7i, R. E., 50
K1Ir.1ro, H., 44Kurokami, T.,IS8, 159
Kush=, L. J., 75
Kw;nctsova, V. S., I49

c.,

Lafferty, H. A., 64. 70

Lakon, G_, lC, I19, 121, 122, 124, 125
Lambou, M. G.l#l, 98, 127, 152, 166, 172
LaPine, L. J., 611

Large,]. R.,9 8
Larter, L. N. I:!,;1 66
Lnughland, D. J-L, 71
Laugltland, J., 71
Leach, Woo, 148
Lee, M. R., 185
Lco;art, C. W., IIS
Lcgnini, C. N., 114
Leroux, D., 75
Lesagf:, P., 117
Leus, F. P., 139
Lewis, D., 157
Libby, W. F., 4, S
Lindemann, E., 152
Lindstrom, E. \V., 36
Lipkin, B. H., 83
Lishkc\ich, M. I., 152
Lee, J. R, 17z
Long, T. E., 161
Loomis, W. E., 179
Lopez, M, 98
Lute, A. M., IS0, 139
Luthra, J
5
Lyncs, F. F., 47

c.,

McAlister, D. P., 38
McC:Ug, J. D., 183
McCalb, A. G., 64
McClelland, T. H., 98
McFarlane, V. H., 94
Machacek,]. E., 45
r.1cHargue, J S., 117
McKee, R., 18
McLean, D. Moo, 54
McLeish, J., 157
McLemore, T. A., 184
MacLeod, A. M., 67, 126, 134

~'-

MacM..a.>ters, M. M., 149, 168, 190

McNeal, X., 38
?>.kRostie, G. P., 66
Madsen, S. 13., 53
l\1anarcsi, A., 84MaIT:lS, F., 124Marrero, J., 98
Martin, J. A., 54
Martin, J N., 70
Martin, S. c., 76
Mason, T. G., 73
Mathur, P. B., 75, 189, 191
Mattson, A. 1\1., 134l\1azl.:ov, F. F., 120
Mehta, M. L., '91
Merrill, L. G., Jr., 191
McrrilI, S., Jr., 183
Milberg, E., 68
Miles, L. E., 4-8
_Miller, E. V., 136
Miller, P. R., 18
Milner, M., 4-0, 44, 48, 49, 65, 72, 149, 18 5
Mirev, N. Too, 83, 95, III, 15:;:
Misonoo, G., 64Moe, N. G:t 3
Monsclisc, ::i. P., 119
Moo~,~. P., li4MooIJaru, M. N., 150
Morrison, B. Y., 89
Moss, E. H., 87
MOlirult, A., 176
Miiller, F. yon, 89
Miiller-Olsen, c., 136, 137
Mucnscher, W. c., 91
Mukherji, D. K., Iz6

Munn, M. T., roo

r.lunn, R. E., 100
Murray, M. D., 188
Murray, M. J., 171
Murv.in, H. F., 179
Musil, A. F., 18
Myers, A., 50, 53, fiq, 70, 76, 77, 106, ,60
Niidvemik, J, 127
Nagel, C. M., r90
Nilijima, Y., 87
Nakatomi, S., 117
Nanda, K. K., Il6
Navashin, Moo, 154. 156
Nnar<wa, N. S., I17
Neljubnw, D., 1I7, II8
Nelson, c., II3
Nelson, M. L., 83
Nelso~ O. E., Jr., 136
Nemec, A., lIS
Nesbitt, L. L., 74, IS:;:
NCllStadt, M. H., 168
Nicholson,
5
Niethammer, A., liS, II8
Nib-n, R. A., xi, IS3, 166, 177
Niles, J. J., 124
Noecker, N. L., 67
Nord, E. c., IIS

c.,

North,

c., 28

Nuccorini, Roo, 116

Nuwe, G. E., 53
21 3

......

Oathout, C H., 71
O'Connor, R. T., 188
O'Donnell, W. W., 152
OgasaW.U;l,

K., 182

Qyam, 0., II5, 117, 149

Ramircz-Sih-a., F. J, 179
Ramstad, P. E., 71, rRoo, M. N., 15'
Rees, B., 37
Rchac1wva, 0., 83
ReilZ., L. P., 74
Reynold~,.E. B., ro
Richards, A. V., 97
Riddell, J. A., '39

160

Rios, Gu=ilez- -see Gonzilcz-Rios, P.

Rippertoll, J. C, 72
Rist, C E., 168
Rirvancn, T., 12{
Roach, J. R., '74Robbins, W. A., 72
Roben, A. L., 46
Roberts, L. \V., '34
Robertson, D. W., % 65, 67, 150
Robertson, F. R., 152
Rodrigo, P. 1\.., 53, 70, q.o
Rodriguez-Colon, F., C}8
Roe, E. I., 8z
RohmC<1cr, E., 133
Romberg, L. D., 192
Romm, H. J., 189
Rose, D. H., 50
Rousseu, N. Brocq-, 151
Royce, R., Iii
Rudolf, P.O., 83
Ruge, U., ITO
RI1S("3, R. A., N
Russell, R. c., 45

Ogden, W.D., 179

Ohga, I., 4, 13,35, 138
Okamura, T., 35, 9+. 150, 151, 152
Olafson, J. H., 148, 17'
Ohen, Milller- -Set Miiller...()!sen, C.
0ppt'Tllnlln, 0., 95
Osterman, E., 61
Owen, E. B., \;
O>:ley, T. A., 49, 65, 67, 159
Pagan, V., 179
Painter, E. P., 74. 152
P:mikkar, M. R., 159
Papavizas, G. C, x,4O,.p
Parker, J., 133
P:lCkinson, A.. II., 138
Paton, R. R., 95
Patrick, S. R., 68
Paul, A.. D., 183
Pearce, J. A., 152
Pedro, A. V. Mil, 53
Peel, A. C, 106
PciItt, G. J., 173
Perch'al, J., 67
Pen'Uhina, N. V., 8j
Petersen, K.. Dorph-, j'fj, lIZ,
PercTliOhn, E., I16
Petiver,3
Peto, F., 15.1
Petty, D. H, 179
Pfahler, P. L, 174
Phillis, E., 73
Pickett, C. A., 185
Pingale, S. V., 16,
Plaut, M., 119, 120
l'ohloudek-l'abrini, R., 152
Pons, W. S., Jr., 152
Poole, H., 1'27
POptso\", A. V., 76
Poptzoff, A., 75
Poncr, R. H., In, 72, 126
Porrer, G. F., 183
Prasad, M., IS{
Predtctshenskaja, A. A., Iii
Pringle, W. J. S., 197
PritchaId, E. W., 50. 53
Puntambekar, S. V., 152

"'C'

Sacks, W., 179

Safono"", Shefer_ -Sa Shefer-Safono,-a,
E.IA.
Sabs, W. W., tOO
Sabta, T., IIB
S..mck, j., 68
Sampicrro, G., 35, 94Sanders, H. G., 53
Saran, A. n., 94
S:trgt:nt, C. S., j8
Sayre, }. D., 34. 66
Schenk, R. V., 126
Schjclderup-Ebbe, T., 3
Schloesing, A. Th., 75
Schmidt, E. W., 75
Schmidt, J. L., 160
Schmidt, W., Il6
Schubert, G. H., 83
Sehwass, R. H., 147
Schweiger, E., I.2..j., 134
Schwcmmle,j., '53
Scott, D. H., 77
Semeniuk, G., 40, +;
Senti, F. R., lQO
Screnlov, G. 1'., t49
Shamel, A. D., 75
Shcherba\;ov, V. G., 182
She:lT, C. 8., 98
SHedd, C. K., 76
Shefer-S..(onova, E. lA., 117
Sherman, G. D., 172
Sherman, W. A., 160
Shirasawa, H., 83
Shishinr. E. D. H. EI-, j6
Shkvarnikov, P. K., 1$3
Sh.hamikov, P., 154Shud, R. W., 123
Sifton, H. 8., fry
Si1.1l, Ramirez- -sa Ramircz-5ilva, F.).
Simak, 111., X, 136, 137, 185
Simpson, D. M., 18, 33, 34. 72, 73

"4

.. , . . .. ,

.-..

. ;:.-

",~.'

: 0.

~N~~. ~Fi -:

...
Singh, B. N., 116
Singh, K. 1Grp:ll, 184
Singh, S. G., 49
Singh, S. L., 49
Singley, M. E., 161
Sloane, 3
Slusanschi, H. t 67. lSZ
Smith, A. J. j"l., IS
Smith, C. L., S9
Smith, E., 13-+

Smith, F. G., 134

Smith, G. M., 174Smith, L., ri, 35, 37. 157. 177
Snyder, G. B., 100

Solo,-je'-", 1\-1. A., S8

SODa\'llC, K... 10.1., 70
Sorenson,}. W., Jr., 160
Sorgcr-DomcniggJ B., 40, 41, 44, (20, 152
Sp:1cth, j. N., lIb
Spencer, G. E. L., 71, 75
Spencer, H. M., 16,25
Srceniv:LS:l.Il, A" 149. 151
Stansbury, M. F., '52
Starooubtsm<l, A. 1., 149
Steinb:mer, C. E., 8]

Steinbauer, G. P., 83. 86, 1 q

Stevens, O. A., 69, 70
Src,enson, F. J., 54
Stewart, F. C, 67
Slolle, A. L., 53
Stone, B. :M., 72
Street, O. E., 120
Srub\)t" H" '53, 154, 156
Subrahmanyan, Y., 189
SulliwIl, J. T., 174-

Suskin, ,'.1., 48, roo

5ycnson., j., 12,!Svihla, R. D., 61

S...~c, G., 66, r6I
Swaminath:m, M., t89

Swanson, C. 0., 64
Takahashi, R., 181
Takematsu, T., 182
Tiliguchi., Y., II6
Tammes. '1'., 36
Tanashe,, G. E., liS
Tarr, S. A. J., 73
Tashiro, S. A., J49
Tatnu.n, E. c., 50
Taufcl, K., 152
Taylor, C. A., 76, 86
Tecson, A. L., 53
Tejima, '1'., 64
Terasalta, Y., 94, 181
Teclsvuori, K., 6g
Tervet, l., 72
Teunisson, D. J., 46
Thal"\lm, A. G., 159
Thomas, A. S., Sg
Thomas, B., +IThompsoll, J. 8., 72
Thor, C.]. B., 192
Throneberry, G. 0., 134
Tieghcm, P. van, 6
Tillotwn, C. R., 78

Todaro, F., (n
Toniolo, L., II9, 124
Toole, E. H., xi, 9, 13, 18,22,37, 53, 68,
72,7-1-, '40, 170
Toole, V. K.,:ri, 13, 37, 53, 69, 72, 170
Tourney, J. W., 711, 83
Towers, B., 73
Tr:mlWein, K., 112
Treccani, C. P., 113
TSchizo\":l., A. "'I., 177
Tskoidze, V., II7
Tuite, J. F., 40, 43
Tukey, H. B., 112, 113, 114
Tupper, M. F., 5
Turcsson, G., 118
Tumer, J. H., 1,3
Turrill, W.ll., 7
Tunle, A. P., 100
U!tec, A.

J., 98

van Doren-ut Doren, C. A. van

van Ticghern-set Tieghcm, P. \":In
Vcnl..:l1ram, C. S., 44

Velhey, c., 126

Verr",t, J. A., 95
Viado, j., JI2
Vib:tr, T., 70
Vlgnes, J. L., 172
Villafane, A. G., 98, 179
Vincent, G., 38, 8)
VlSW:1l1:ltha, T., 1!l9
\"Oil. Degen-se: Dcgell, A. Yon
yon Miiller-uc Miillcr, F. von

Wach, A., JI9

Waggoner, Ii. D., 54, lSi
Wahlen, F. '1'., 37
Wakele)', P. c., 31, 78, 79, 82
Walkden, H. H., 76, 192
Wallace, H. A. H., 45
WaUace,~I.

AL, 196

Wallen, V. R., 111

WaUee, A. D., 136
Walster, H. L., 65, 67
Ward, l-I. S., 149
Ward, H. S., Jr., 28
Wark, D. c., 47
Wasserman,]., lIZ
Watanabe, S., 54
Webster, C. c., gll
Weibull, G., 33, 54, 62 t60

Weise, R. t 119

Wci5cnfeld, M., liZ

Weiss, M. G., 36
Wellon, F. A., 67
Went, F. W., 101
Wentz,]. B., 36
\harton, M. J., 125, 120
Whecler, W. A., 69
White, D. G., 98
White, G. D., 172
White,]., 63

"'5

69,

Wl1ite, O. A'I 5
Whyper, K., 63. 64

Wrighf, R. C., S9
Wyttenbach, E., 152

Wilbur, D. A" tT:1

Williams, M., 68
Williams, R. W. M., 88
Wilson, ."r.. K.. 8]

Y:magisawa., T., 133

Young, G. Y., 65

Windisch, 120
Winstanley, J., 115

Winlers, I-l F., 98

Zabclin2, Z. K., 18a

Zadwiew, B. J.t II1

\~olJc:n~eber.lI.

W., H
\\oodbndge, M. E., IllS
Wright, E. C.

LONGEVITY

7.ek:ny, L. t 152. 168

Barton-,s

Zimmerman, P. W" 178

216