Principios y Practicas de Almacenamiento de Semillas

PRINCIPLES AND PRACTICES
OF SEED STORAGE
By Oren L. Justice and Louis N. Bass
Agriculture Handbook No. 506
On January 24, 1978, four USDA agencies—Agricultural Research

Service (ARS), Cooperative State Research Service (CSRS), Extension
Service (ES), and the National Agricultural Library (NAL)—merged to
become a new organization, the Science and Education Administration
(SEA), U.S. Department of Agriculture.
This publication was prepared by the Science and Education Admin-

istration's Federal Research staff, which was formerly the Agricultural
Research Service.
Library of Congress Catalog Card No. 78-600015
Washington, D.C. Issued April 1978

CONTENTS
PRINCIPLES AND PRACTICES
OF SEED STORAGE
By Oren L. Justice1 and Louis N. Bass2
INTRODUCTION
Purpose and Significance of Safe Seed Storage
The principal purpose of storing seeds of economic plants is to
preserve planting stocks from one season until the next. Prehistoric
man learned the necessity of this practice and developed methods of
storing small quantities of seeds for his future use. As agriculture
developed, man expanded his knowledge regarding both the require-
ments of seed for maintenance of viability and methods of providing
suitable storage conditions. In 1832, Aug. Pyr. de Candolle cf France
included a chapter on seed preservation in his book "Physiologie
Végétale." He pointed out that the vitality of seeds would be prolonged
if stored under conditions to protect them from heat, moisture, and
oxygen. About the same time, other authors suggested the use of
tar-coated wooden boxes and iron tanks capable of being sealed with
stopcocks for drawing off the seeds as desired.
Although the storage of seed stocks for planting the fellowing season
remains the most important reason for storing seeds, farmers and
seedsmen have found it advantageous to carry over seeds for 2 or more
years. This practice results in accumulating supplies desired genetic
stocks for use in years following periods of low production. Many kinds
of seeds—mostly vegetable, flower, and forage seeds—move rather
freely in world commerce. Many of these seed lots are not used the first
year after production.
With increased knowledge and technology of plant, genetics and plant
breeding, the necessity for longtime storage of small quantities of the
various cultivars becomes apparent. Facilities for storing genetic stocks
now exist at Fort Collins , Colo., Hiratsuka, Upan, Braunschweig,
Germany, Bari, Italy, Izmir, Turkey, and other locations. Research
workers can obtain genetic stocks considered useful in their breeding
programa from these facilities. The principies to be discussed here
AGRICULTURE HANDBOOK 506, U.S. DEPT. OF AGRICULTURE
apply equally to all seed storage. Longtime storage of genetic stocks is

not the principal objective of this handbook.
No statistics are available indicating either the actual losses sustained
from substandard storage conditions or the savings effected by opti-
mum storage conditions. However, the importante of suitabie storage
becomes apparent when one considera the alternatives. Failure to use
available information about seed storage could greatly handicap the
Nation's agricuItural program as follows: (1) Agriculture in the warm,
humid, subtropical, and tropical regions would be less efficient , (2) the
plant breeder would be greatly handicapped by not maintaining genetic
stocks, and (3) international commerce in seeds would be only a small
fraction of its present economic value.
The Seed, a Fragile, Living Organism

In his poem "A Package of Seeds," Edgar A. Guest expressed very
well the theme of this section. When Guest had purchased packages of
seeds, he had thought of them only as seeds, But on one occasion when
he made such a purchase, it flashed through his mirad that he had
purchased "a miracle of life," a dime's worth of power that no man can
create and a dime's worth of mystery, destiny, and fate. In working
with seeds, especially in harvesting, cleaning, handling, storage, and
transportation, it is essential to keep in mind at all times that incide the
seed is a dormant miniature plant awaiting the opportunity to continue
its growth.
Basically the seed is made up of the embryo or miniature plant, the
endosperm and other food reserves, and protective coverings consisting
of the seedcoat and, in some cases, accessory structures (fig. 1). The
embryo is more protected in the resting seed of corn than in bean. In
corn and other grains the major food reserves are stored as endosperm,
whereas in bean they are stored in the two cotyledons or seed leaves,
which also serve as photosynthetic organs for the young plant. Other
structures affecting seed storage are the seedcoat for both corn and
bean and the hilum and micropyle of bean, which may permit the entry
or exit of moisture. Under some conditions the hilum may become soft
or damaged, allowing fungi to enter the seed.
The planting units in several plant families are fruits, botanically
speaking, rather than true seeds. In the grass family the seed or fruit is
called a grain or caryopsis, in the buckwheat and sunflower families an
achene, and in the mint family a nutlet. Some kinds of seeds possess
additional structures, such as glumes, bracts, spines, and hairs, which
aid in protecting the seed from injury, birds, and rodents. In this
publication, all these structures that normally serve as the planting unit
are referred to as seeds.
PRINCIPLES AND PRACTICES OF SEED STORAGE
The orientation of the embryo within the seed and the nature of the
protective coverings are important attributes of any seed that is sub-
FIGURE 1.—External and internal structures of corn (A) and bean (B and C) seeds.
jected to the numerous mechanical and handling processes from harvest

to planting. For example, a grass seed has its embryonic root tip near
the base of the grain. When the glumes are removed during harvesting,
as in rye, timothy, and wheat, the root tip is only protected by a thin
membrane, which is very vulnerable to physical damage and attack by
storage molds. However, seeds of many grasses, such as barley, oats,
rice, and some cultivars of sorghum, are much better protected since
the enclosing glumes usually remain attached.
The relatively large seeds of bean and soybean have only a moder-
ately thick seedcoat. The plumule, or embryonic stem, is fairly well
developed in the resting seed and lies between two cotyledons, or seed
leaves. Also, the radicle, or embryonic root, has practically no protec-
tion except that provided by the seedcoat. Thus, a bean seed is unu-
sually vulnerable to breakage, especially when dry and roughly treated.
The radicle, plumule, or cotyledons can be damaged. Velvetbean, in the
same plant family as bean and of similar structure, is less vulnerable te
damage from harvesting, handling, and processing because of a slightly
different orientation of the plumule and radicle.
From Harvest to Sowing

Beginning with harvest, seed lots usually pass through a series of
processes necessary for immediate or future use. They include har-
vesting, drying, cleaning, grading or sizing, transporting, and storing.
Not all commercial seed lots are subjected to all these processes, but the
manner in which seeds are passed through each process can affect their
storage potential.
Each seed faces an uncertain future. It may be harvested too early,
too late, too wet, or too dry. The seed's future can be affected by being
harvested so immature that it cannot germínate, or if it germinates, it
may produce a weak seedling. The moisture content of the seed may be
so high that heating will occur before the seed is dried to a safe moisture
content; it may be damaged during drying, or it may be so dry that it
suffers impaction damage on handling. The potential danger during the
drying process is.exposure of very wet seeds to high temperatures or to
temperatures in the critical range for a long time.
In the cleaning and the grading or sizing processes the seed may again
be subjected to impaction damage, which may be caused by low mois-
ture content in the case of bean and many other croes harvested under
arid or semiarid conditions. Radish seed may be damaged severely by
impaction if its moisture content is too high during threshing. High
moisture makes the seeds difficult to remove from their pods.
When transported to market, whether a few miles or across the
ocean, the seed lot is subject to storage under various adverse condi-
tions—on the truck, in the railway car, on the dock while partially or
entirely exposed to the elements, or in the hold of ships. A seed lot for
use in the following year or years would normally be placed in a
conventional storage facility, which may or may not be conducive to
maintaining vigor and viability. Because of the high capital investment
and cost of maintaining desirable seed storage facilities, many seed lots
are stored under unfavorable conditions.
During processing and shipment, grain for industrial uses is fre-
quently moved through suction-type conveyors at great speed and
force, impacting with metal at curves and bends, and falling great
distances. This results in considerable breakage and other damage. The
amount of this damage can usually be calculated and is tolerated, since
the grain will be processed for food, feed, oil, or other industrial uses.
Grain or other types of seeds to be planted should not be subjected to
such treatments. Damage from impaction and improper drying or
storage may or may not be obvious on casual observation. Seeds may
receive internal fractures from impaction, moisture stress, or heat
stress without corresponding damage to the surface.
Decline and Death of Seeds

This publication is concerned primarily with the varlous factors
affecting the life and death of seeds in storage. Decline in vigor and
death of seeds can be considered from two aspects: (1) Loss of viability
or death of a seed lot, i.e., a small or large quantity of seed, or (2) death
of an indlvidual seed. Ordinarily people are interested in the viability of
a quantity of seed as this is what they most likely will plant. They want
to know whether the germination capacity of the seed lot is sufficiently
high to produce a satisfactory stand of plants and, if so, the required
rate of planting to produce a stand. The germination percentage of a
seed lot is the proporton of individual seeds capable of producing
normal plants. For this reason the decline in vigor and eventual death of
an individual seed should be considered.
Seeds of most crop species are mature when they attain maximum
dry weight. Most seeds are physiologically mature at this point, but
there are exceptions. When physiologically mature, the seed possesses
its greatest vigor. From this point on, it gradually loses vigor and
eventually dies. The rate of decline is conditioned by several factors,
some of which man can manipulate. Although he can prolong the
deterioration process, he has not been able to stop it.
Since death of a seed is gradual, it is virtually impossible to determine
when life ceases. Not all the life processes proceed at the same rate in
different seeds of a given seed lot. When seeds are stored under various
conditions, these life processes can terminate in individual seeds at
different times. For example, certain enzymes remain functional for a

time after a seed is dead.
Woodstock (1973) expressed biological deterioration of seeds as
follows: "In cellular tercos there is no sharp line of demarcation between
life and death in seeds. Rather, seed death is a gradual and cumulative
procesa as more and more cells die until certain critical parts of the seed
become unable to perform their essential function."
Moore (1955) summarized the transition from seed maturity to death
as follows: The so-called vigor of life is at its greatest leve! when the
seed first matures. Alter maturity, however, that life becomes less and
leas vigorous as the never-ceasing aging process moves onward toward
death; but even long before death the seed becomes questionable or
worthless for planting purpose, especially under field conditions that
are not highly favorable for germination and seedling development.
"This transition of life at its fullest, to death with the ever associated
loss of vigor as intermediate steps, may take place slowly or rapidly,
depending opon the kinds of seed, the extent and location of mechanical
injury, the moisture and temperature conditions to which the seeds are
subjected in the field, during harvest, and in storage, and the important
factor of time during which the seeds are subjected to these conditions.
It is Chis set of conditions that determine the 'true planting age' of the
seed. Seed may thus be surprisingly old, or dead, at a few days or weeks
of age, or distinctly young many years after maturity."
Scope of Publication
Publications on seed storage began to appear in the second quarter of
the 19th century. At least three or four were in French and German in
the 1830's. From about 1840 to 1875 the publications per decade were
slightly lesa than in the 1830's but generally greater in depth. From
about 1875 until today the published reports have become so numerous
that it is literally impossible for one person to review the literature on
seed storage. There seems to have been a great proliferation of papera
from about 1890 to 1910, when more and more were published in
English. In addition to gaining practical knowledge en seed storage,
investigators became interested in determining the longevity of seeds.
Some testad the viability of dry seeds taken from old herbarium
specimen ; others set up longtime experiments by burying seeds in the
soil in such a manner that they could be recovered for future testing.
Owen (195g and Barton (1961) have described these experiments.
We have assembled information here that we hope will be useful to
those interested in storing seeds of vegetables, fiowers, and field crops.
Included is information on the principies of seed storage and their
practical application. Seeds of forest trees, fruit trees, and weeds are
not thoroughly discussed. For information on longtime storage of seeds
for retention of germ plasm, see James (1967, 1972) and Ito (1972).
Many changes occur in seeds during storage. The principal changes
considered in this publication are three quality factors—germination
capacity, vigor, and yield potential. Since much information exists on
chromosome aberrations and mutagenic changes in aged seeds as well as
biochemical changes related to enzymes, enzymatic action, and metab-
olites of these processes, these subjects are treated only briefly.
SEED FACTORS THAT AFFECT STORAGE LIFE

The seed's characteristics, inherent conditions resulting from natural
causes, and man's treatment and handling affect the seed's storage life.
The storage potential of an individual seed is influenced by the following
10 characteristics and conditions and possibly by others.
Genetic Effects
Variation Among Species
Man learned early that seeds of some plant species would survive a
given set of storage conditions longer-than others. This information is so
common today that it may appear unnecessary to mention it here;
however, a few examples are documented.
Essentially all known cases of seeds surviving for a century or more
belong to species with hard impermeable seedcoats. Harrington (1972)
in his list of species with longevity records of 10 or more years included
four genera in this group: Albizia 147 years, Cassia 158 years, Goodia
105 years, and Trifolium 100 years. Ali belong to the Leguminosae, a
plant family known for producing hard seeds. An exception is a report
by Aufhammer and Simon (1957) that barley, which had been sealed in
glass tubes and placed in the foundation stones of a building in Nurem-
burg, Germany, showed 12-percent germination after 123 years. Most
species known or estimated to have survived for over 500 years are hard
seeded; for example, species of Canna (Anonymous, 1968), Lotus
(Ohga, 1923), and Lupinus (Porsild et al., 1967).
In contrast to these leguminous seeds, the seed of peanut, also in the
legume family, is relatively short lived. Seeds of economic species with
a short storage life include lettuce, onion, parsnip, and peanut. Of the
cereals, barley and oats usually have the greatest potential for satis-
factory storage, rye probably the least, and corn (maize) and wheat are
intermediate (Haferkamp et al., 1953). Haferkamp et al. (1953) found
that three lots of 32-year-old barley germinated 96, 80, and 72 percent,
whereas seeds of four lots of oats stored under the same conditions
germinated 84, 66, 55, and 6 percent. MacKay and Tonkin (1967) found
barley to have about the same storage capacity as wheat and consider-
ably leas than that of oats.
Variation Among Cultivars
Inheritance of seed longevity is not limited to species. Several studies
show that the principles prevailing at the species level are also effective
at the cultivar level, at least for some crop species. Several cultivan of
the same species that were compared for seed longevity differed
significantly. Shands et al. (1967) reported that seeds of the cultivar
Oderbrucker barley had greater resistance to germination loss in
storage than other cultivars. Toole and Toole (1954) found that the
cultivar Black Valentine bean had a longer lifespan in storage than
'Brittle Wax.' James et al. (1967) found significant differences in the
storage of various cultivan of bean, cucumber, peas, sweet corn, and
watermelon. Their review of relevant literature indicates that the work
of Lindstrom (1942), Sahadevan and Rao (1947), and Haber (1950)
consistently supported the probability of dominante of longevity. In a
study of eight soybean cultivars, Burgess (1938) found germination
variations between 21 and 99 percent after 4 years compared with 95 to
99 percent after 5 months' storage. In this study the differences among
cultivan did not show up until the third and fourth years of storage.
Results of studies on inbred limes of corn by Neal and Davis (1956)
suggest that some inbred lines maintain viability better than others
when stored at low temperatures. Lindstrom(1942, 1943) found that
some inbred lines of corn germinated 90 percent after 12 years' storage
at room conditions in Iowa, whereas all seeds of other inbreds were
completely dead. As indicated previously, he also found that increased
longevity in corn inbreds appeared to be dominant; however, he also
pointed out that inheritance of longevity is not simple. He believed it is
possible to introduce genes for longevity finto various fines of corn by
backcrossing. Weiss and Wentz (1937) reported short seed longevity
and low vigor of corn to be associated with luteus 2 and luteus 4 genes
found on chromosome 10. Other luteus genes on the same chromosome
did not decrease storage life of the seeds. The authors were unable to
explain this apparent contradiction.
Generalizations
Not all seed species, cultivars, or individual seeds within a genetic
group are destined to survive for the same period of time under a
specified set of conditions. A lot or sample of seeds does not die at one
time, but the individual seeds making up the lot or sample die over a
period of time. In referring to storage lile, lifespan, period of viability,
or storage potential, we mean the length of time required for a certain
percentage of the seeds to die or conversely for a certain percentage to
PRINCIPIES AND PRACTICES OF SEED STORAGE
live. Different percentages have been used for different purposes

(Roberts, 1972).
In summary, the seeds of different plant species vary widely in their
lifespan under identical or favorable storage conditions. Only a few
studies have been made to determine the inheritance of long storage life
among cultivars and most of them have pertained to corn (maize). This
might be a worthwhile subject for further study. In planning farm or
commercial storage the expected storage life of the species should be
considered. In experimental studies every effort should be made to use
homozygous cultivars, Unes, or strains, and studies should be conducted
under strictly controlled conditions.
Preharvest Effects
Austin (in Roberts, 1972) reviewed the literature on the effects on
viability of the environment during seed development. Included were
such factors as temperature, photoperiod, mineral nutrition, rainfall,
and soil moisture. However, he did not relate these studies to seed
storability. In fact, few, if any, of the studies we have reviewed were
designed to test the relative effects of preharvest factors on seed
storability.
How Seeds Are Affected
Desirable candidates for storage are matare seeds of normal size and
appearance and relatively free of mechanical injuries and storage
micro-organisms. They should not have been subjected to extreme
temperature and moisture conditions during maturation and harvest.
Thus, any preharvest environmental factor that influences these seed
qualities also affects the storability of seed lots. About 95 percent of the
total dry weight of a seed is storage material, which will be used by the
germinating seed and seedling until it can produce its own photosyn-
thetic and nutrient absorbing machinery (Pollock, 1961). It is readily
seen that an immature seed, a seed with an unbalanced chemical
composition, or one mechanically damaged, permitting early entry of
micro-organisms, would be at a disadvantage in storage.
Barton (1941) found that seeds of high initial viability resisted un-
favorable storage conditions better than similar seeds of low initial
viability. Working with Austrian winter pea, McKee and Musil (1948)
showed that seed germinating 94 percent did not deteriorate under upen
storage at ambient temperatures and humidities at Beltsville, Md., and
four locations in the Southeastern States as rapidly as similar seed that
germinated 75.5 percent at the beginning of the experiment. Brett
(1952a, 19526) reported that seeds of grasses, legumes, vegetables, and
root crops grown in England with high initial germinations deteriorated
leas rapidly than similar seeds of low initial germinations. MacKay and
Tonkin (1967) cheeked Brett's work by using large numbers of samples

stored under laboratory conditions at Cambridge, England. They con-
firmed Brett's results for seeds of grasses, root crops, and vegetables
but not for legumes. They concluded that it was unlikely that hand seeds
in legumes accounted for this behavior.
Bunch (1958) adequately summarizes this discussion: "Previous
treatment of the seed, in the field, during harvest, or in previous
storage may have affected the seed in such a way as to lessen its chances
of storing well. In warm and humid climates the seed may deterlorate
on the plant before harvesting. Immature seeds and mechanically
damaged seed are especially liable to lose viability rapidly in storage."
Effect of Provenance
Very little information is available on the effect of provenance, or
location of production, on seed vitality and apparently none relating
provenance directly to storability of seeds. MacKay and Tonkin (1967)
compared the time required for seeds of four forage species grown in
different countries to deteriorate to 80-percent germination. The results
were as follows: Red clover seed grown in Canada required 4 years
compared with 3 years for English- and New Zealand-grown seed;
perennial ryegrass seed grown in Ireland and New Zealand required 4
years; American-grown meadow fescue required nearly 7 years com-
pared with 6 years for Danish-grown seed; and Irish-grown crested
dogtail seed required 6 years compared with 3 years for New Zealand-
grown seed. These data may show significant differences among loca-
tions of production; however, the seeds were not stored under con-
trolled conditions and sources of seeds were not predetermined; rather,
the data were limited to the seed receipts at the Official Seed Testing
Station at Cambridge, England. Thus, no definite conclusion can be
drawn at this time as to the effect of provenance on storability of seeds.
Effect of Weather
The most obvious preharvest factor affecting seed viability and
storability is weather, especially seasonal changes. Farmers and seeds-
men alike know the perils and risks of excessive moisture and freezing
temperatures during the later stages of seed maturation and postmat-
uration stand in the field. Some documentary evidence follows:
Dillman and Toole (1937) stored seed of four fiax cultivars grown
under irrigation in Califorria in 1929 and 1930. The 1930 seed "showed
marked weather injury and a low test weight (36.4 to 45.0 pounds per
bushel)." The germination of four cultlvars after 6 years of storage of
the 1930 crop was 1, 4, 0, and 9 percent, whereas the corresponding
germination of the 1929 crop of the same cultivars was 94, 86, 87, ard 94
percent.
MacKay and Tonkin (1S67) showed positive correlations between

weather conditions during ripening and harvesting of barley, oats, and
wheat and the number of years required for the seed to deteriorate to
either below 80-percent or below 50-percent germination. The data
were obtained from large numbers of samples grown in England over a
period of 26 years. Average hours of sunshine and inches of rainfall at
Cambridge were used as indices of weather.
Results obtained on seeds of numerous species stored in an unheated
building up to 33 years at ambient conditions near Lind, Wash., led
Haferkamp et al. (1953) to conclude that healthy matured seed stored
unthreshed and harvested in dry weather is a prerequisite for long
storage life. Riddell and Gries (1956) suggested that variations in
growth of spring wheat from seed of different ages were related to
temperature of maturation rather than to age of seed or storage
conditions. Harrington and Thompson (1952) found that the region
where lettuce seed was grown significantly affected its germination at
between 24° and 30° C. Highly significant positive correlations were
established between germination percentages and the temperature 10
to 30 days preceding harvest.
Moss et al. (1972) found that preharvest rains can cause wheat to
germinate in the ear accompanied by increased activity of a-amylase,
which digests some of the starch. Fortunately there appears to be some
variation among cultivars for activity, suggesting the possibility of
selecting or breeding varieties with resistance to this characteristic.
Early sustained freezes in the Corn Belt while seed corn still in the
field has a high moisture content can cause serious damage. The degree
of damage is affected by the temperature reached, duration of low
temperatures, and moisture content of the seed. Effects of the damage
may become obvious soon after a hand, sustained freeze. Some seeds in
such a lot or entire seed lots exposed to leas severe freezes may be
damaged only to the extent that the effects are detectable after storage.
Because of the seriousness of the problem and its importance, corn
subjected to freezing has been investigated to a considerable extent.
Some other crop species have been studied also but on a smaller scale.
For additional information, see page 30.
Seed Structure and Composition

The presence or absence of glumes (lemma and palea) in grasses is a
well-known example of a seed structure that influences lifespan. Ha-
ferkamp et al. (1953) found that aged seeds of three cultivars of barley
and Red Winter Speltz wheat with hulla retained viability heder than
seeds of the same harvests that had been threshed and stored. Other
cultivan of barley and wheat, as well oats and rye, showed the same
trend but to a less extent. Huila, chaff, or both had an inhibitory effect
on the growth of mold, suggesting that the increased lifespan of cereal

seeds was due to suppression of mold growth by the glumes during
storage. Lakon (1954) showed that oat and timothy seeds had a longer
lifespan when stored with the glumes intact compared with similarly
stored machine-hulled caryopses. Seeds of both species behaved simi-
larly. From Lakon's studies the hulled seeds appeared to be damaged
mechanically during removal of the hulls.
Working with seeds from six districts in Sweden, Esbo (1954) re-
ported that the viability of hulled timothy seeds declined 16 percent
during the first year, whereas seeds with the hulls intact showed no
significant loss in viability until the third year. Also, he found that field
emergence of unhulled seed was 6 to 14 percent higher than emergence
of hulled seed. Average germination of six seed lots after approxi-
mately 2 years was 78 percent for unhulled and 40 percent for hulled
seed.
Goff (1890) found that hulled timothy seeds, on sale or stored in
Wisconsin, did not germinate as well the first year nor after storage up
to 10 years as did unhulled seeds. The differences were as follows:
The smaller difference for seed stored 10 years may have been because
the unhulled seed declined in viability after 8 years of storage. Although
Goff used samples from commercial seed lots in his studies, he sug-
gested that the reduced germinability of hulled seeds could not have
been caused by thresher injury "as all seed of either sort that showed
the least indication of injury were rejected from the trials."
Stevens (1935) stored unhulled and hulled timothy seeds under the
same conditions for 11 years. During this period the germination of
unhulled seed declined from 98 to 52 percent, whereas the hulled seed
declined from 97 to 16 percent.
Since the work of Goff and Stevens, several workers have shown that
many kinds of seeds suffer interna] damage not always apparent with-
out dissection and special tests (Esbo, 1954). Esbo pointed out that
when the loosely attached palea and lemma are detached from the
caryopsis, the latter is vulnerable to mechanical injury. He deflnitely
associated the rapid decline in viability of hulled seeds with mechanical
injury.
Roberts (1972) discussed the protection offered several shapes of seed
to mechanical damage during harvesting, handling, and processing.
Generally small seeds escape injury, whereas large seeds are more
likely to be extensively damaged. Size, arrangement of essential seed

structures, and seed composition are contributing factors. Bean and
lima bean are examples of seeds highly susceptible to damage. The large
cotyledons and location of the embryo axil represent a structure that
will tolerate only low level impacto. Although corra is, relatively large,
flat seed, its strong pericarp, strong adherence of parta, and location of
embryo offer some protection, yet injuries can be expected to the root
tip, since it is only moderately protected. According to Roberts (1972),
spherical seeds usually give more protection than flat or irregularly
shaped seeds. Of course, the spherical shape is mediated by other
characteristics, such as relative exposure of vital parts (root and shoot
axes), thickness and strength of seedcoat, and brittleness of all seed
parts at the time of impact.
Justice (1950) referred to the relatively high incidence of abnorrnal
seedlings in onion, as did Clark (1948). Roberts (1972) observed that the
embryonic root tipa of onion extend beyond the seed, a condition
conducive to mechanical injury.
Available literature does not relate seed composition or biochemistry
to lifespan. Conceivably some relationships may exist, but more definite
information is needed. Chemical, physical, and nutritive changes during
storage are discussed by Zeleny (in Anderson and Alcock, 1954) and
biochemical indices of deteriorating seeds by Abdul-Balri and Anderson
(in Kozlowski, 1972).
Hard Seeds
Much has been written about hard seeds, or seeds that are imperme-
able to water within the usual germination period, but little information
is directly related to seed storage.
Most crop species in the legume or pea family (Leguminosae) produce
hard seeds of varying percentages. Harrington (1916) indicated that the
important legumes cultivated in the United States probably produce
varying percentages of impermeable seeds except the peanut. Many of
the current croe species can develop hard impermeable seeds when
produced and stored under certain environmental conditions. Although
hard seeds are usually associated with species of the legume family,
they may also be produced by species in other plant families. For
example, hard seeds are frequently found in hollyhock and okra and
infrequently in cotton in the mallow family (Malvaceae), catnip in the
mint family (Labiatae), cranesbill in the geranium family (Geraniaceae),
canna in the Cannaceae, Ipomoea, Convolvulus , and Cuscuta in the
morningglory family (Convolvulaceae), and Indian Lotus in the Nym-
phaeaceae.
Possibly impermeable seeds are produced by several species in addi-
tional families, especially by the native wild planta. In fact, Passerini
(1933) explored this possibility, but his data fail to distinguish between
retarded germination caused by impermeable seeds and physiological
dormancy, which may be unrelated to seedcoat effects.
Most of the literature does not state clearly the developmental stage
in which the different legume seeds become hard. According to Lowig, 4
fully ripened seeds of white clover contained slightly more hard seeds
than less mature seeds. Lute (1928) found that the percentage of
impermeable seeds of alfalfa increased with maturity. Harrington
( 1915) indicated that 90 percent or more of well-matured seeds of alsike
clover, red clover, white clover, and white sweetclover were hard. It
appears evident that Harrington referred to the condition of seeds
before storage Martin ( 1944) found that most lots of red clover seed
obtained from 13 growers in Iowa contained a considerable number of
hard seeds, varying from O to 35 percent. Also, Martin (1945) reported
that sweetclover seed lots coming from the harvester contained, on the
average, more than 50 percent of hard seeds. Under certain environ-
mental conditions the hard seed content of some lots increases during
storage.
Helgeson (1932) showed that practically all slightly immature sweet-
clover seeds were permeable when harvested but were made imperme-
able by drying them over calcium chloride for 7 days or without the
calciurn chloride if kept in dry storage. Impermeability or seed hardness
was prevented by storing the seeds in a moist, cold room. Generally a
warm, dry atmosphere is conducive to hard seed formation and a cool,
moist atmosphere favors a low hard seed population. in many species,
hard impermeable seeds have a longer lifespan than permeable seeds.
This is an obvious advantage to the survival of the species but causes
problema in cultural practices. The current tendency with annual crops
is to plant at a rate that will produce the number of plants desired in a
given area, as uniform germination is necessary te produce uniformity
of crop, permitting "once-over harvesting." Hard impermeable seeds
that germínate at some future date have little planting value under
these circumstances. On the other hand, storage of seed at intermediate
or high relative humidity to minimize the percentage of hard seeds in a
lot contributes to more rapid deterioration of germination capacity,
vigor, or both.
Varying percentages of hard seed of most species become permeable
during overwinter storage in the temperate zone. Softening depends
largely on temperature and relative humidity. Harrington (1916) con-
ducted extensive studies on the hard seed conditions in small-seeded
legumes. He concluded that nearly all hard seeds of alsike clover, white
clover, and sweetclover in dry storage remained impermeable for at
least 2 or 3 years. Impermeable red clover seeds gradually became
permeable, but 30 to 65 percent of the hard seeds remained imperme-
able for 4 years or longer. Most alfalfa and hairy vetch seeds became
permeable within 2 years.
Martin (1945) in Iowa found that up to 24 percent of the hard seeds of
sweetclover softened when stored at temperatures around freezing, but
softening was nil at 15°-30° C. This confirmed the observations of
Stevens and Long (1926). Presumably the seeds were stored at ambient
humidities in Iowa. Martin (1944) reported that hard seeds of red clover
from 13 sources softened to a maximum of 33 percent, depending on
source, when stored dry in a laboratory at Ames, Iowa. Two seed lots
changed from a relatively low grade, based on germination percentages,
to a fair grade during a short period of storage. According to Lute
(1928), 50 percent of the hard seeds of alfalfa became permeable within
3 1/2 years when stored in Colorado.
Researchers have consistently reported great differences in the per-
centages of hard seeds among species and among seed lots of the same
species (Whitcomb, 1931; Esdorn and Stuetz, 1932). Esdorn and Stuetz
(1932) reported that hard seeds of lupines, alfalfa, and red clover
softened rapidly under the suitable temperature-humidity conditions,
whereas softening in hard seeds of alsike clover and white clover
proceeded very slowly.
Much has been written about the structure of leguminous seeds in
relation to water absorption. A few references are cited here. Seedcoat
or testa (Rees, 1911; Watson, 1948), cuticle (White, 1909), hilum
(Hyde, 1954), strophiole (Martin and Watt, 1944), and a general paper
by Coe and Martin (1920). The anatomy of some other crop species that
produce hard seeds is discussed by Simpson et al. (1940) for cotton,
Reeves (1936) for cotton and other malvaceous plants, and Gaertner
(1950) for Cuscuta (Convolvulaceae).
The following genera and number of species (in parentheses) of field
crops, vegetables, herbs, and flowers can be expected to produce hard
seeds. They are included in the "Rules for Testing Seeds" (Association
of Official Seed Analysts, 1970).
Some snap bean cultivars produce hard seeds, although most do not.
Hard seeds in snap bean are undesirable. Among cultivars containing
hard seeds are Top Notch and Golden Wax (Barton, 1966a), Decatur
and Blue Lake (Nutile and Nutile, 1947), White-Seeded Kentucky
Wonder (Harrington, 1949), and Green Savage (Joubert, 1954).
Lowig 5 listed 41 cultivara of bean, mostly of German origin, with hard
seeds varying between O and 97 percent. Thirteen of these showed hard
seeds occurring at 50 percent or aboye, and 21 cultivars showed hard
seeds exceeding 25 percent. These figures must be used with caution
since Lowig determined hard seed percentages after storage over
calcium chioride.
Researchers in Puerto Rico (Anonymous, 1942a) found that drying
beans with calcium chloride was unsatisfactory because of the hard
seeds produced. Nutile and Nutile (1947), Harrington (1949), and Bar-
ton (1966a) demonstrated the effect of low relative humidity on hard
seed formation in bean. For example, Harrington stored White-Seeded
Kentucky Wonder beans with an initial hard seed content of 33.5-per-
cent and 8.3-percent moisture content. After 60 days of storage at
10-percent relative humidity, 74.4 percent of the seeds were hard,
whereas samples stored at 65-percent relative humidity for the same
time contained no hard seeds.
It is obvious that seed of certain snap bean cultivars can change
readily from permeable to impermeable depending on the relative
humidity of the storage area. Such is not so with seed of other species.
Seed of several small-seeded legume species became hard when stored
at a low relative humidity and remained hard when transferred to a high
relative humidity because the valve-type closure to the hilum permitted
moisture to escape from but not enter the seed. When impermeable
seeds of most species became permeable, they remained so regardless
of the relative humidity where they were subsequently stored.
In some trade practices, hard seeds are regarded as viable. Several
scientific papers attest to the fact that not all hard seeds are viable, for
example, Goss (1926) and Toole (1939) for hairy vetch. Crosier and
Patrick (1952) found the viability of hard and nonhard hairy vetch seeds
to be similar during the first 5 years of storage, but after 17 years both
were among the dead ones. Also, dead hard seeds have been reported in
okra.
Nutile and Nutile (1947) and Harrington (1949) made suggestions for
storing bean seed that may have recome hard with age. Nutile and
Nutile recommended that such lots not be stored in artificially heated
buildings. Harrington recommended that storing such seeds at about
21° C and a relative humidity of 50 percent or possibly a little higher
should prevent hard seed formation except possibly for very low
percentages in the most susceptible cultivars.
Seed Maturity
Relationship of Maturity to Storability
Several scientists have considered seed maturity as being that stage
when maximum dry weight has been attained (Harrington, 1972; Rob-
erts, 1972). Since many crop species fiower, produce, and mature seeds
over a period of several days or even weeks, it is important to know at
what stage of maturity the seeds should be harvested. Much of the
research relating to maturity has been oriented toward production and
harvesting goals without continuing into storage studies. Such studies
have marginal relevante to the relationship of seed maturity and
lifespan in storage. Since healthy, mature, plump seeds generally store
better than immature seeds, any studies that show the relationship
between seeds of different maturity levels and germination, vigor, field
emergente, or yield relate indirectly to storage and lifespan. Many such
studies have been discussed by Harrington (1972) and Austin (in
Roberts, 1972).
Grass Seeds
Several studies on the relationship between seed maturity and stora-
bility or longevity have been concerned with grasses. Both the flower-
ing and seed maturity characteristics of the grass plant may extend
over several days depending on crop species. This results in harvesting
seeds of varying stages of maturity from recently fertilized ovules to
mature, plump seeds. Many of the light grass seeds are removed in the
cleaning processes; however, many are not, especially those in the
premilk and milk stages.
McAlister (1943) collected seeds of three species of Agropyron, three
species of Bromus. Elymus glaucus, and Stipa viridula in the premilk,
milk, dough, and mature stages and stored them at between 15° and 23°
AGRICULTURE HANDBOOK 506, U.S, DEPT. OF AGRICULTURE
C at moisture contents of 7-9 percent. Evaluation tests made alter 4

and 5 months showed that the seeds harvested in the premilk and milk
stages were generally inferior as to both viability and longevity to seeds
harvested in the dough and matare stages; the only exceptions were
that viability of seeds of Bromus marginatus and B. polyanthus
harvested in the premilk and milk stages was equal to that of seeds
harvested in the dough and mature stages.
Similar resulta and conclusions have been reported by numerous
workers, including Hermann and Hermann (1939) for Agropyron cris-
tatum, Griffith and Harrison (1954) for Phataris arundinacea, Esbo
(1959) for Phleum pratense, Bass (1965) for Poa pratensis, and Jorgen-
sen (1969) for Festuca pratensis.
Other Field Crop Seeds
Only limited work has been done on seed maturity and storage of
large seeds. However, studies have shown in many instantes that
immature or partially filled seeds are inferior to mature seeds in
viability and vigor. These include Dimmock (1947) for corn, Blackstone
et al. (1954) for peanut, Inoue and Suzuki (1962) for bean, and Turner
and Ferguson (1972) for cotton. Justice and Whitehead (1942) found
immature velvetbean seed to be inferior in viability to mature seeds.
The germination for vine sublots that could be compared were mature
seeds 73 percent and immature seeds 10 percent. Storage period varied
with sublots from 3 to 7 months depending on the date of harvest.
Vegetable Seeds
Two Japanese workers reported some relevant information on the
effect of maturity on vegetable seed storage. Eguchi and Yamada (1958)
atorad seeds of 11 kinds of vegetables (cabbage, carrot, Chinese cab-
bage, cucumber, edible burdock, eggplant, Japanese radish, pumpkin,
tomato, watermelon, and Welsh onion), which they liad harvested at 3-
to 7-day intervals giving 4 to 6 stages of maturity for 2 to 3 years. Their
English summary indicates that eight kinds showed marked loases in
viability of immature seeds compared with mature seeds. However,
Chinese cabbage, pumpkin, and tomato showed little or no difference in
longevity with different maturities.
Pollock (1961) pointed out that maturity studies of carrot are compli-
cated by the fact that at least three orders of flowers, or umbels, are
produced and each order will flower and produce seeds over a period of
time. Harvesting all umbela at the same time will result in both mature
and immature seeds. He reported that the leves of maturity affects both
germination and vigor.
Austin (in Roberts, 1972) harvested carrot seeds in England at four
dates from September 5 through October 5 and classified each harvest
into four size groups. As the harvest season progressed, the weight of
seeds declined slightly, but the germination increased by 5 percent,
which probably was not significant. As the size of seed increased, the
mean seed weight increased correspondingly. The germination in-
creased with increased seed size for the first three weight classes (1.00
to 1.75 mm per seed) and for each harvest date, but it decreased by 4
percent for the largest and heaviest seeds (1.75 to 2.00 mm per seed).
These studies suggest that immature carrot seeds do not germinate as
well as mature seeds. Based on studies with seeds of other crop species,
the longevity of immature seeds would be apt to be less than that of
mature seeds when both are stored under identical or similar conditions.
Germination response of seeds removed from mature and immature
squash fruits differed presumably because of seed maturity. Young
(1949) and Holmes (1953) collected mature and immature fruits of
butternut squash and stored them for periods up to about 7 months. At
the beginning of his experiment, Young found seeds from mature and
immature fruits to weigh 7.78 and 5.32 gm per 100 seeds, with average
germination of 90.8 and 19.0 percent, respectively. Four months later,
both mature and immature seed had gained weight by 0.34 and 0.91 gm
and germination had increased to 98.4 and 67.2 percent, respectively.
At the beginning of the experiment, seeds from only 7 out of 25
immature fruits germinated, whereas a11 immature fruits produced
germinable seeds at the conclusion of the experiment. This indicates
that the seeds of some immature fruits either were too immature to
germinate when harvested or were dormant.
After 7 months of storage of butternut squash, Holmes ( 1953) found
the weights per 100 seeds to be 10.9 gm for mature and 8.9 gm for
immature seeds. Germination at this time was 97 percent for mature
and 77 for immature seeds. Holmes (1953) also found that the ratio of
fruit weight of typical butternut squashes to seed weight was rather
consistent.
Seed Size
No referentes have been located relating seed size or weight to
lifespan through storage experiment. On the other hand, numerous
studies have shown the superiority of heavy, mature seeds over light,
immature seeds in germination, vigor, and yield tests. Relatively few
exceptions have been noted. Comprehensive reviews are provided by
Black (1959) and Austin (in Roberts, 1972). Austin (1972) discussed seed
maturity and seed size jointly, possibly because seed size is determined
or mediated to some extent by maturity. His treatment lends credence
to the view that the same or similar vagaries besetting immature seeds
in storage also affect small seeds, at least to some extent.
Seed Dormancy
Relationship of Dormancy to Storage
Some or all seeds of several crop species are dorrnant at harvest. A
major problem facing seed technologists when testing stored seed for
germination is how to overcome dormancy. Dormancy of freshly
harvested seeds may be found in practically all groups or classes of plants,
whether crops or native piaras, including grasses, cereals, clovers and
other small-seeded legumes, large-seeded legumes including peanut,
cucurbits, vegetables, flowers, trees, and weeds. In some seeds, dor-
mancy is caused by (1) seed structures, as seedcoats, bracts, glumes,
pericarp, and membranes, which limit the exchange of water and gases,
(2) physiology cf the embryo, (3) germination inhibitors or other blocks,
or (4) a combination of these factors.
Storage can affect dormancy in many instances. In most crop species,
dormancy is dissipated within a few to several montes when the seed is
stored at ambient temperatures and relative humidities or under con-
trolled atmospheres provided the temperature is held aboye freezing.
Seed physiologists know that the best method of maintaining dormancy
in seeds is to store them at subfreezing temperatures.
Dissipation of Dormancy
Owen (1956) and Koller (1972) reviewed pertinent literature on the
existente of seed dormancy and its dissipation with time. Only a few
relevant studies are mentioned here. Larson et al. (1936) found that
dormancy in immature seeds of barley, oats, rye, and wheat persisted
longer than in mature seeds when stored at 0° to —35° C, but that
dormancy varied greatly among cultivare of a given crop species. The
duration of dormancy could be increased by lowering the temperature.
In a study of seeds of different cultivars of barley, oats, and sorghum,
Brown et al. (1948) found that incipient dormancy was overcome in most
cultivars within 1 to 6 montes when stored at 40°. At 2.2°, dormancy in
barley and oats persisted for 3 years.
Dore (1955) and Roberts (1972) discussed dormancy in rice. In Dores
(1955) studies of 21 cultivars, dormancy was lost between the 7th and
11th weeks after harvest except for the cultivar Penifun, which was
dormant for only 5 weeks. Roberts (1972) found the dormancy of rice
cultivars to vary between practically none in the cultivar Tai Chu 65 to
over 100 days (100 days for 50 percent of the seeds to break dormancy)
in Masalaci. Although dormancy among rice cultivan differed mark-
edly, the pattern of viability lose for all cultivars was the same.
The situation with grasses is similar to that for cereals. In many
native American grasses used for range planting, seed dormancy is
much more intense than in cereals. Winchester (1954) found that
Panicum maximum var. Trechoglume seed increased in germination

from 11 percent when fresh to 46 percent after storage for 25 months
and seed of Cenchrus ciliaris (buffelgrass) from 3 to 79 percent within
13 months.
Dormancy in Florida runner peanut seed was overcome by Hull (1937)
when he stored it at 20°-25° and 40° C, whereas dormancy in some
samples persisted for 2 years at low temperature storage conditions.
Recently harvested cottonseed can be dormant. Christidis (1955) re-
ported that some cottonseed samples grown in Greece remained dor-
mant up to 5 months. Justice (1956) found that the only effective method
of breaking dormancy in seeds of Cyperus rotundus was to place them
on a moist substratum at 40° for 3 to 6 weeks.
Dormancy and Lifespan
In considering the relationship of seed dormancy to storage, a second
aren of interest is whether dormancy increases the lifespan of seeds.
Roberts (1972) reviewed pertinent literature on the subject and cor-
rectly concluded that available evidence is not sufficient to establish
even a casual relationship. He detailed and analyzed some of his own
experimental work with rice and concluded that his work did not
support the existente of a relationship between dormancy and longev-
ity. This does not exclude the possibility that the lifespan of dormant
seeds of native or wild plants buried in the soil under natural conditions
my be longer than the lifespan of nondormant seeds.
Toole and Toole (1953) found that imbibed dormant seeds of lettuce
did not decay in 105 days, whereas companion samples placed in beakers
with a relative humidity of 85 to 90 percent were dead; the seeds of one
cultivar remained alive for 42 days. This suggested to the authors that
dormancy of the imbibed seeds either held in check or suppressed the
life processes that lead to seed deterioration.
Existing evidence is not sufficient to demonstrate a positive relation-
ship between seed dormancy and lifespan. However, dormant seeds in
storage undergo changes, some of which lead to breaking of dormancy
and others to inducing dormancy. Some of these changes are affected by
storage conditions.
Moisture Content
The moisture content of seeds during storage is no doubt the most
influential factor affecting their longevity. It is important to harvest
mature, relatively dry seeds or to reduce the moisture content of
high-moisture seeds soon after harvest. Bass (1953) found that the losa
of viability of freshly harvested Kentucky bluegrass seed was corre-
lated with the moisture content of the seed and length of time held at a
given temperature. Seed with 54-percent moisture held at 30° C for 45
hours lost 20-percent germination, but seed with 44-percent moisture

withstood 45° for 36 hours with no apparent loss of viability. Seed with
22- and 11-percent moisture content showed essentially no loss in
viability at 50° for over 45 hours.
McNeal and York (1964) studied the effects of drying sorghum seed
on viability. They concluded that sorghum to be used for seed should be
harvested when the moisture content is about 20 percent or less and
dried promptly to about 11-percent moisture content. The drying tem-
perature should not exceed 43.3° C for very moist seed and not over
54.4° for seed with a relatively low moisture content.
Haferkamp et al. (1953) reported reduced germination of 27-year-old
wheat compared with slightly older wheat. They explained this on the
basis that the June rainfail in 1923 (year of harvest of 23-year-old
wheat) was over five times the average. The authors believed that high
moisture content of the seeds during development greatly reduced seed
viability and longevity. Regarding moisture during seed development,
Mcllrath et al. (1963) made an interesting observation on developing
tomato seeds. Although the moisture content of the placenta and
pericarp of tomato fruits 15 days old or older never dropped below 94
percent, the seeds became dehydrated to approximately 50-percent
moisture content as the organs developed and matured.
Another way in which initial seed moisture content affects storage lile
of seeds is through its effect on mediating damage by threshing and
processing machinery and handling. Although it is very important to
reduce seed moisture content to a safe level for storage, it is also
important to be aware of possible adverse effects of low moisture
content. Very dry seeds are susceptible to mechanical breakage and
related injuries. Such damage may result in physical breakage or
fracturing of essential seed parts, make the seed vulnerable to fungal
attack, and reduce storage potential. For more information, see the
section on "Effects of Storage Environment on Seed Longevity."
Mechanical Damage
Awareness of Seed Damage
As harvesting and threshing machinery and the combine carne into
general usage, damage to seeds and grain increased accordingly. Some
of the early work on seed injury related to the mutilation of corn
(Brown, 1922), seedcoat injury of barley and wheat (Hurd, 1921),
broken pericarp of corn (Myers, 1924), and mechanical injury to soy-
beans (Oathout, 1928). Research on seed injury was expended greatly
in the 1930's, with the advent and use of the combine thresher. Some
seed species or groups have received greater attention than others
depending on their vulnerability to damage by machinery and handling.
Two important characteristics that affect the degree of damage are seed
structure and resistance of the seed to removal from the pod, as in
legumes and crucifers, or from the mother plant, as in the grasses.
Problems arising from these characteristies are affected by moisture
content of seeds and pods, degree of maturity, and possibly other
factors.
Beattie and Boswell (1939) and Moore (in Roberts, 1972) showed that
damaged seeds do not store as well as intact seeds and that fungi enter
the seed through cracks in the seedcoat. In discussing the storage of
seeds of the clovers and medies, Brett (1952a) stated, "From the results
of experiments at Cambridge [England], and elsewhere, it is clear that
scarifled seed loses its ability to germinate at a significantly greater rate
than does unscarified seed when stored under the same conditions."
Natural deterioration in spinach seed has demonstrated that the
percentage of abnormal seedlings increases with time and the percent-
age of normal seedlings decreases. Thus, seedling abnormalities result
not only from mechanical damage but also from natural aging. Accord-
ing to Moore (in Roberts, 1972), small and hidden injuries in seeds,
including bruises, may not cause immediate loss in vitality, but they can
become increasingly critical with aging of the seed. Moore (1972)
pointed out that injuries to vital organs, ie., various parts of the
embryo, become more serious with age than injuries to nonembryonic
tissues.
Vulnerability of Some Seeds to Damage
The amount of fiber in the bean pod, which varies with cultivars,
affects the degree of mechanical damage to the seeds through its effect
on ease of threshing. Apparently this problem was first studied by
Harter (1930) of California, then continued and expanded by his col-
leagues Borthwick (1932) and Bainer and Borthwick (1934). A com-
prehensive review of the subject by the Asgrow Seed Co. (Associated
Seed Growers, Inc., 1949) also included resulta of its own extensive
research studies. This type of research has been continued since the
1950's, especially at the U.S. Department of Agriculture, Beltsville,
Md. (Toole, 1950; Toole et al., 1951; Toole and Toole, 1960) and at
Michigan State University (Perry and Hall, 1960, Dorrell; Dorrell and
6
Adams, 1969). These studies showed that threshing or combining

produces breaks, cracks, bruises, and abrasions in seeds, which in turn
result in abnormal seedlings of questionable planting value. These
injuries were reproduced by controlled mechanical shock to the seeds
and were greater with reduced seed moisture.
Toole (1950) attempted to compare the germinability of slightly

injured seeds with seeds producing apparently uninjured seedlings of
bean stored for 3, 6, and 9 months. Owing to unpredicted changes
during storage, such as (1) decrease in apparently uninjured seedlings,
(2) corresponding increase in slightly injured seedlings, and (3) increase
in abnormal seedlings, no firm conclusions could be drawn. However,
the author suspected that under unfavorable conditions slight mechani-
cal injuries could predispose the seed to increased deterioration com-
pared with uninjured seed, even though the preliminary study did not
confirm this suspicion.
Response of Seeds to Storage Conditions
There is reliable evidence that damaged seeds of small-seeded leg-
umes do not survive as long as nondamaged seeds. Battle (1948) found
that all alfalfa seeds that had been scarified with sandpaper and stored
for 14 years were dead, whereas unscarified seeds showed 23-percent
germination. Similar results were reported by Graber (1922), Stevens
(1935), and Brett (1952a). In storage experiments, Oathout (1928) and
Mamicpic and Caldwell (1963) demonstrated that damaged soybean
seeds lose their viability earlier than nondamaged seeds. Blackstone et
al. (1954) reported similar results for peanut.
Information on the retention of viability of injured cereals in storage
is extremely limited. In a 17-year experiment Russell (1958) compared
lots of wheat seed having cracked seedcoats with lots of sound seed. At
the beginning of the experiment the damaged seed germinated 4
percent less than the sound seed. This difference gradually increased to
12 percent, where it remained for 9 years of storage. The germination
of the two groups differed slightly during the last 4 years of storage.
Possibly the sound seed declined rapidly in viability after 10 to 12 years.
Hurd (1921), Kuperman (1950), and Wortman and Rinke (1951) showed
that broker. cereal seedcoats provide easy access for microflora to enter
the seed. Broken or cracked seedcoats also enhance embryo damage by
chemical treatments, including chemicals used as disinfectants (Bonvi-
cini, 1951). Both fungí and chemical damage reduce the keeping quality
of stored seeds.
Possible Involvement of Genetics
Efforts have been made to determine whether there are genetic
differences among bean cultivan in their tolerance to mechanical in-
jury. Atkin (1958) studied 3 crops of 18 cultivan and concluded that
important fairly constant differences existed. Generally the colored-
seeded cultivan had greater resistance to mechanical injury than
white-seeded cultivars. Nevertheless one white-seeded cultivar
(Streamliner) showed some resistance to mechanical injury. This led the
author to believe that a resistant white-seeded cultivar could be devel-
oped. On the other hand, Dorrell 7 found that tolerante to seed injury of
navy bean is a complex trait controlled by numerous genes. Population
averages showed no evidence of dominante in the F 1 and F 2 generations
and genetic variante appeared to be primarily additive. Thus, from this
study there is little encouragement that cultivars highly resistant to
mechanical injury can be developed through a breeding program.
It is obvious from available information that mechanical injury to
seeds not only reduces production of normal seedlings but also de-
creases the storage potential of damaged seeds that apparently would
have produced normal seedlings prior to storage. Many studies have
been made on seed injury resulting from mechanical harvesting,
threshing, combining, and handling, but relatively few of these studies
have been concerned with storage.
Vigor
The vigor of seeds at the time of storage is an important factor that
affects their storage life. Vigor and viability cannot always be dif-
ferentiated in storage experiments, especially in seed lots that are
rapidly deteriorating. Irrespective of this problem, several workers
have demonstrated that seed iota that were rapidly declining contained
some seeds low in vigor and other seeds that still might be vigorous.
This progressive weakening with age continued until all the seeds
became nonviable.
Obviously new, vigorous seed lots possess a greater storage potential
than older lots that may be approaching rapid deterioration. General-
ized curves of the decline in vigor and viability of a seed lot with time
are shown in figure 2. The viability curve gradually decreases, followed
by a sharp decline and finally a gradual loas. The loss in seed vigor
essentially parallels that of viability but at lower levels. The rate at
which seeds will decline in vigor or viability depends on several factors,
including genetic constitution of the species or cultivar, condition of the
seed, storage conditions, uniformity of seed lot, and storage molds if the
relative humidity permits their growth. Among the research workers
who have shown that vigorous seeds reach a point when deterioration
proceeds more rapidly than during the early period of storage are
Patrick (1936) for Chewings fescue seed, Christidis (1954) for cotton-
seed, and Burns (1957) for blue lupine seed.
The decline in vigor and viability of seeds is sometimes illustrated by
a sigmoid survival curve. The survival curve for dry seeds stored under
favorable environmental conditions can be dissected into three distinct
parts. The first represents the period when the seed is vigorous and
decline in the life functions has proceeded slowly. Eventually this stage
FIGURE 2.—Decline of viability and vigor with time of a typical seed lot. (Courtesy
of J. F. Harrington.)
ends at a survival level of 90-75 percent, and deterioration then

proceeds very rapidly. After deterioration has proceeded to a survival
level of 25 to 10 percent, it slows again and continues slowly until all
seeds are dead. The curves for vigor and viability are very similar,
except that loss of vigor precedes loss of viability.
EFFECTS OF STORAGE ENVIRONMENT ON SEED

LONGEVITY
Temperature
Temperatures Above Freezing
Storage temperature and seed moisture content are the most impor-
tant factors affecting seed longevity, with seed moisture content usually
more influential than temperature. Owing to the intricate relationship
between storage temperature and seed moisture content, neither one
can be discussed separately in its entirety. Nevertheless temperature
and seed moisture are discussed here separately to a limited extent to
indicate the relative importante of each, whereas the interaction of the
two is discussed more fully under "Interrelationship of Temperature,
Seed Moisture Content, and Storage Life."
Within limits the storage life of seeds of vegetables, flowers, and field
tropa decreases the temperature increases. Exceptions include cer-
tain short-lived seeds. Groves (1917) described the relationship between
temperature and lifespan of wheat seeds by a formula. Roberts (1972)
presented formulas that he believed described the relationship of tem-
perature and moisture content to the period of seed viability of certain

crop species. He developed nomographs using the formulas for barley,
broadbean, rice, and wheat. The accuracy, limitations, and usefulness
of these formulas and nomographs are still to be determined.
Harrington (1960d) proposed his so-called Thumb Rules, which re-
lated seed moisture and temperature to the seed's lifespan. His (1972)
rules stated that the lile of the seed is halved (1) for each 5° C increase
in seed storage temperature and (2) for each 1-percent increase in seed
moisture content. Harrington cautioned that, based on current knowl-
edge, the first role should not be used for storage temperatures below 0°
or aboye 50°. He also pointed out that the two rules apply independ-
ently, and the effects of temperature and moisture are additive. Since
this procedure gives only rough estimates of storage potential under
stipulated conditions, it must be used with caution.
San Pedro (1936) stored seeds of vegetable species in containers with
calcium chloride at four temperatures and determined the percentages
of germination after 7.1, 9.2, 11.3, and 19.5 months. His results for the
11-month storage period (table 1) show that as temperature was in-
creased, even in a dry atmosphere, germination decreased. His ayer-
ages are mediated by the stage of deterioration of each crop species
during the storage period. For example, cabbage declined in germina-
tion by 19 percent between storage at 0° and 28° C, whereas pechay and
sitao declined only 2 and 3 percent, respectively, under the same
storage conditions. The seeds of radish were dead at all storage tem-
peratures after 14.5 months.
Barton (1941) stored seeds of six vegetable species at different
temperatures and relative humidities for 372 days. At 35-percent rela-
TABLE 1. —Effect of various temperatures on germination of several

crop seeds after storage with calcium chloride for 11 months'
tive humidity the average germination for lettuce and onion, two
species with short storage lives, was 61, 44, 31, and 9 percent, respec-
tively, at 5°, 10°, 20°, and 30° C, whereas comparative germination
results for tomato and flax were 96, 92, 88, and 85 percent.
Barton (1966b) also stored pyrethrum seeds at various temperatures
in sealed and open containers for up to 15 years. Table 2 ,shows that
temperature markedly affects germination. It also shows that the effect
of temperature is influenced by seed moisture. content. In sealed
storage, seed moisture content remains constant, but in open storage it
changes with changes in the relative humidity of the storage ares,.
Therefore adequately dried seeds in sealed containers usually live
longer at a given temperature than similar seeds in open containers.
The effect of temperature on the storage of soybean seed was
reported by Toole and Toole (1946). They stored seeds of the cultivan
Mammoth Yellow and Otootan at five temperatures and three moisture
contents for 10 years. The months required to lose germination at each
temperature-moisture combination are shown in table 3. Not all seeds
were dead at the favorable storage conditions. In fact, there was no
significant losa in germination at the end of the 10-year storage period at
five storage conditions for Mammoth Yellow and at five conditions for
Otootan. The data for 18.1-percent moisture (Mammoth Yellow) and
17.9 percent (Otootan) show the reduction in seed germination or
lifespan as temperature is increased, or alternatively the increased
germination or lifespan as temperature is decreased. The range in
lifespan is from 3 months at 30° C to 72 months at 2° for Mammoth
Yellow and from 1 month at 30° to 72 months at 2° for Otootan. Similar
data could be compiled for other levels of germination, for example 50
or 80 percent.
TABLE 2. —Effect of different temperatures on germination of

pyrethrum seeds in sealed and open storage after various periods'
TABLE 3.—Effect of various temperatures on duration of seed germi-

nation of 2 soybean cultivars at 3 seed moisture contents 11
Small field plantings were made of 10-year-old seeds that had shown
no loss in germination and of seeds grown the previous year. The
authors found no apparent difference in vigor of growth for either
cultivar when the known genetic variation in Mammoth Yellow was
considered. Many seedling abnormalities were observed in both Mam-
moth Yellow and Otootan cultivars grown from seeds artificially dried
to 5.4- and 5.2-percent moisture content, respectively. These abnor-
malities were in seedlings produced from seed stored for only 3 months,
especially from those seeds held at the lower temperatures. Few
abnormal seedlings were produced from seed stored at 300 C, but they
were increasingly numerous at successively lower storage tempera-
tures. The abnormalities of Mammoth Yellow Viere much more frequent
and more pronounced than those of Otootan. Tests made on seeds
stored for longer periods showed approximately the same proportion of
abnormalities.
Some research has been done to determine the effects of high tem-
peratures on seeds. In general, seed viability and vigor are reduced as
temperature is increased, as time of exposure to high temperatures is
increased, and as molture content of the seed is increased. Damage is
diminished at a given temperature as the seed moisture content is
decreased. Thus, Waggoner (1917) found the following relationship in
radish seeds:
AGRICULTURE HANDBOOK 506, U:S. DEPT. OF AGRICULTURE
Harrington and Crocker (1918), Smith and Gane (1938), and Evans
(1957b) showed that low moisture content Kentucky bluegrass, Chew-
ings fescue, and perennial ryegrass seeds can be heated to 100° C or
higher without immediate loss of viability. However, stress tests of
heated seeds showed that vigor decreased before the effects on seed
viability were manifested. White (1909) significantly reduced the per-
cent germination of barley, oats, and wheat by exposing air-dry seeds to
dry heat at 99°-100° for one-half hour. Greater decreases were obtained
by heating for 41/2 hours. Seeds of these three species plus corn (maize)
and rye failed to germinate when the temperature was increased to
122° or higher for 1 hour. The consideration of temperature, time of
temperature exposure, and moisture content of the seeds is essential
when drying seeds for storage. For further information, see the section
on "How Seeds Are Dried."
Temperatures Below Freezing

As background for their experimental results from freezing seeds at
-183°C, Brown and Escombe (1897-98)reviewed at least five papers on
low temperature treatments. They referred to papers by C. de Candolle
and R. Pictet published as early as 1879 and 1885. They frote seeds at
-80° for 2-6 hours, and in 1884 they exposed seeds to -100° for 4 days
without adversely affecting germination. Brown and Escombe (1897-98)
exposed seeds of 12 species with moisture contents of 10-12 percent to
-183° for 110 consecutive hours without damage.
White (1909) observed severe injury to seeds of apple, hemp, lobelia,
parsley, and parsnip after exposure to -200° C for 11/2 days, slight
injury to seeds of bean, mustard, pea, radish, sunflower, and turnip,
and no injury to seeds of cress and Ricinius camogdensis. The moisture
content was not given. She subjected seeds of barley, corn (maize), oats,
rye, and wheat to -200° for 2-3 days without any decrease in germina-
don.
Lipman and Lewis (1934) subjected seeds of 19 species to -250° C for
60 days and evaluated the seeds by greenhouse tests. They found no
significant difference between the frozen seeds and unfrozen controls.
Lipman (1936) exposed seeds of barley, corn, sweetclover, and vetch to
-273.1° just shy of absolute zero, with no impairment of viability,
whereas Becquerel (1953) dried seeds of alfalfa, clover, petunia, and
tobacco and then held them at -272.9° for 2 hours without impairment
of viability.
The superiority of subfreezing temperatures for seed storage com-
pared with higher temperatures has been well established for many
kinds of seeds. Documented experimental evidente dates back to 1928
(Dorph-Petersen, 1928) and possibly earlier. Some of the literature on
seed storage at subfreezing temperatures is summarized in table 4.
TABLE 4. —Sampling of literature on seed storage at subfreezing temperatures
TABLE 4. —Sampling of literature on seed storage at subfreezing temperatures—Continued
1 18-yr data unpublished but supplied by authors.
Weibull (1952, 1955) found seeds of a few species that were not
benefited by a subfreezing storage temperature. A few other examples
have been reported. Roberts (1972) analyzed some of them and con-
cluded that, in some instances, statements were made without consid-
ering the moisture content of the seed. He believed that low tempera-
ture storage is beneficia' to maintaining seed viability, but that there
may be a few exceptions. This position appears to be supported by the
literature.
Early fall freezes while seeds and grain are still in the field have
prompted several studies on the deleterious effects of freezing seed and
grain at high moisture contents. Kiesselbach and Ratcliff (1918, 1920)
studied freezing injury to seed corn because it occurred in the fall with
considerable frequency in Nebraska. They found that death of seed corn
from freezing was directly related to the moisture content of the kernel
and to the duration of exposure to low temperatures. Immature kernels
with a higher moisture content than mature kernels were more sus-
ceptible to freezing injury. All seeds were dead when exposed for 24
hours at the following temperatures and moisture contents:
However, all seeds retained their viability for at least 3 months when
stored at —13.3° to —15.6° C and 12- to 14-percent moisture content.
McRostie (1939) and Rossman (1949) compiled considerable data on
freezing injury of corn seed that essentially confirm the findings of
Kiesselbach and Ratcliff (1918, 1920). All agree fairly well on the factors
causing freeze injury, although their results, when comparable, vary
somewhat in detail as would be expected. According to McRostie (1939),
5-day fluctuating temperatures of —17.8° to —23.3° C, —17.8° to —26.1°,
and 0° to —26.1° caused more damage to corn seed than a constant
exposure to the lowest temperature of any of these regimes. On the
other hand, Rossman (1949) indicated that repeated freezing and thaw-
ing were less injurious than continuous freezing when the total time was
the same. He found similar responses to freezing temperatures by the
two hybrid cultivars he used. Likewise, Kiesselbach and Ratcliff (1918,
1920) found that dent corn and flint corn responded similarly.
In storage experimenta with frost-damaged sweet corn seed, Barton
(1960b) found no significant loss of germination in five lots (four
cultivars) over 9 years when storescaled at —5° C with moisture contents
below 13 percent. In open storage at 30° or at —5° in a moisture-satu-
rated atmosphere, the seed decreased in viability by about the same
amounts over 7 years.
Apparently seeds of barley, rye, and wheat are nearly as sensitiva as

corn to cold injury at high moisture contenta. Agena (1911) found a
rapid increase in injury in these species when the grain was stored at
-6° C and moisture contents aboye 22 percent. Under these conditions,
injury was evident after 1 day of storage and increased with time. He
found that moist grain can be held best for short periods between O and
-6° but that fungi soon appear under these conditions. Freezing injury
was greatest at the radicle tip and secreting anea of the scutellum.
Wheat was frozen in liquid nitrogen (bp —195.8° C) for 2 minutes at
moisture contenta of approximately 19 to 25 percent by Lockett and
Luyet (1951). Entire seeds with 19.4-percent moisture content dropped
10 percent in germination because of freezing. Corresponding loases in
germination with increase in moisture content of entire seeds were as
follows:
Imbibed decorticated seeds had a considerably lower moisture content

with less germination loss because of freezing than entire seeds imbibed
for the same time. The moisture content of imbibed embryos was much
greater than that of either entire or decorticated seeds.
Freezing injury to sorghum seed is affected by moisture content of
the grain, temperature, and duration of freezing treatment (Robbins
and Porter, 1946; Carlson and Atkins, 1960; Kantor and Webster.
1967). In addition, Carlson and Atkins (1960) showed that the genotype
of the cultivar affects tolerance to freezing injury. Data of Robbins and
Porter (1946) aleo suggest genetic effects on freezing tolerance; how-
ever, a definite statement to this effect cannot be made since the
complete history of each seed lot used is not given. Kantor and Webster
(1967) used two cultivara in their research but were unable to draw a
conclusion with respect to genotype effects. In view of these reports on
sorghum seeds and the reports of Kiesselbach and Ratcliff (1918, 1920)
and of Rossman (1949) on corn, the genetic effect on freeze resistance in
cereals remains unresolved.
Seed Moisture Content and Relative Humidity

Direct Effect of Moisture on Seed Deterioration
Barton (1961) regarded moisture content of utmost importance in
seed deterioration. Seed deterioration increases as moisture content is
increased.
Boswell et al. (1940) illustrated the effects of relative humidity and
consequently moisture content of the seeds on their storage life. They

stored 10 kinds of vegetable seeds at 2 temperatures, 3 relative humid-
ities per temperature, and under warehouse conditions for 251 days.
Samples were tested on O day, every 10 days thereafter for 90 days, and
after 110 and 251 days of storage. Since 10-, 30-, 50-, 70-, and 90-day
tests were not made on peanuts, data were extrapolated to obtain these
results for averages.
The averages for 12 test dates times 10 species, or 120 tests per
storage condition, are as follows: At 26.6° C, with relative humidities of
78, 66, and 44 percent, the respective average germinations were 55.4,
78.9 and 81.3 percent. At 10°, with relative humidities of 81, 66, and 51
percent, the respective average germinations were 80.7, 82.6, and 82.6
percent. In some respects these resulta do not reveal the true situations
for the following reasons: (1) The storage period was not long enough to
measure deterioration of long-storing species, such as tomato or several
other species at the more favorable storage conditions, e.g., 10° and
51-percent relative humidity. (2) Averaging species such as onion, with
a short storage life, with tomato cancels out the extremes of both
species. For example, the percent germinations of these two species on
the 251st day of storage at 26.6° and relative humidities of 78, 66, and 44
percent were, respectively, onion, 0.1, 37.4, and 72.9 percent and
tomato, 68.1, 85.0, and 91.8 percent.
Since seed deterioration is affected by moisture content, it is impor-
tant to know what factors affect water absorption and retention as well
as their effects. Obviously the thickness, structure, and chemical com-
position of the seedcoat affect the rate of water absorption and reten-
tion by seeds; in hard seeds the seedcoat restricts total water uptake. Of
the various seed constituents, proteins are most hygroscopic (readily
taking up and holding moisture), carbohydrates are slightly less so, and
the lipids are hydrophobic (lacking an affinity for water). Thus, seeds
containing relatively high percentages of carbohydrates, proteins, or
both, such as rice, other grains, and soybeans, can have moisture
contents of about 13-15 percent at 25° C and 75-percent relative
humidity, whereas cottonseed, flaxseed, and peanuts, which are rich in
oil, would have moisture contenta of approximately 9-11 percent at the
same temperature and relative humidity (Barton, 1941). Barton found
that seeds of flax, lettuce, onion, peanut, pine, and tomato showed
differential water absorption and each species retained its same position
relative to other species in all her experimenta.
Conceivably a sample of seeds containing both living and dead seeds
might take up a different amount of water than a sample of only live
seeds or, alternatively, all dead seeds. In studies of this phenomenon by
Atkins (1909), Heinrich (1913), Barton (1941), and Simpson and Miller
(1944), all agreed that under storage conditions the total moisture
uptake and retention do not differ between living and dead seeds. Seeds
used included bean, cotton, flax, lettuce, onion, peanut, perennial
ryegrass, pine, and sweetpea.
One of Harrington's (1960d) so-called Thumb Rules of drying seed
stated that for each 1-percent reduction in seed moisture content, the
time the seed can be stored without seriously affecting germination is
approximately doubled. Harrington (1972) also indicated that this rule
applies when seed moisture content is between 5 and 14 percent.
Roberts (1972) developed formulas relating temperature and seed
moisture content to the storage life of seeds and indicated that Har-
rington's (1960, 1972) rule agreed with his formulas. However, Roberts
(1972) pointed out that possibly there are inaccuracies in his own
formulas for both seed moisture content and temperature and
suggested that the same may be true for Harrington's (1960, 1972) rule. On
the other hand, both Harrington (1960, 1972) and Roberts (1972) stated
that their procedures give approximate results only.
The doubling of the lifespan of seeds by decreasing the moisture
content by 1 percent shows vividly the effects of slight moisture
changes, especially at the critical point. In sealed storage the expected
life of vegetable seeds with an 8.0-percent moisture content could be
doubled by removing 1.0-percent moisture before sealing. On the other
hand, certain possible pitfalls should be understood and considered
when working near the critical moisture level. Some of these are—
(1) The degree of confidence by which an aliquot of seed tested for
moisture represents the lot or bulk.
(2) The degree of accuracy in grinding, drying, and weighing.
Usually the air-oven methods are accurate to no more than ±0.1
percent.
(3) The possible variation in seed moisture content among different
lots or crops. This can result from differences in chemical composition of
the seed caused by different climatic conditions and cultural practices
during seed development and maturation.
(4) The hysteresis effect, a phenomenon when at a given relative
humidity the equilibrium moisture content of grain and seeds may not
always be the same. When seeds lose water and reach equilibrium at
any stated relative humidity, the equilibrium moisture content of the
seeds may be higher than if dry seeds are allowed to gain moisture to
reach equilibrium at the same relative humidity. This means that seeds
with a moisture equilibrium value of about 12.5 percent at a relative
humidity of 60 percent could possibly have two moisture equilibrium
values depending on whether the original seed moisture content was
greater or less than 12.5 percent. Hlynka and Robinson (1954) showed a
difference of about 4.5 percent at a relative humidity of 40 percent for
wheat but no difference at 90-percent relative humidity. Hubbard et al.
(1957) found adsorption equilibrium values for corn and wheat to be

approximately 1.6 percent higher than desorption values at 33- and
22-percent relative humidity, respectively.
The most common method of determining the moisture content of
seeds is to hect them in a forced air oven at a given temperature for a
specified time or until constant weight is obtained. The losa of water
represents the moisture content of the seeds. The percentage of mois-
ture can be calculated (1) by the wet weight method—the amount of
water lost is divided by the initial weight of the sample, and (2) by the
dry weight method—the amount of water lost is divided by the dry
weight of the sample alter drying. Since the original weight in (2) is
divided by a smaller number than in (1), the answer will be greater for
(2). Thus, in equivalent situations dry weight percentages will be
slightly greater than wet weight percentages.
Percentages of moisture based on dry weight are frequently used in
research and percentages based on wet weight are usually usad for
commercial purposes. Seedsmen and seed testing laboratories (Assoc.
Off. Seed Anal., Internad Seed Testing Assoc., and Soc. Com. Seed
Tech.) use the wet weight method to determine seed moisture content.
Many U.S. scientists also use ths method. However, if the basis for
calculating the percentage of moisture is not stated in the literature, it
is risky to assume that a specific procedure was followed.
Relationship Between Relative Humidity and Seed Moisture

Content
As previously mentioned, the moisture percentages of seeds in equi-
librium with a specific relative humidity vary with crop species. At any
temperature air will hold a given amount of water in the form of vapor.
When the air contains all the moisture it will hold, it is said to be
saturated, which is equivalent to the dewpoint or 100-percent relative
humidity. Ordinarily the sir is not saturated but contains only a fraction
of the amount of water it would hold if saturated. Relative humidity
reflects the amount of moisture actually in the alas a percentage of the
amount of moisture that the air is capable of holding at the same
temperature. Thus, relative humidity, expressed as a percentage, is
determined as follows: The amount of moisture in the air is divided by
the amount of moisture the air is capable of holding at the same
temperature and multiplied by 100. Warm air can hold more water than
cool air. Thus, if the amount of water in the air is held constant and the
temperature is increased, the relative humidity will be decreased.
Conversely, if the temperature of the air is lowered, the relative
humidity will be increased.
The importante of seed moisture content in retarding seed de-
terioration cannot be overemphasized. Under all storage conditions the
moisture content of the seed will come to equilibrium with the sur-
rounding air if given enough time. In fact, equilibrium is reached
between the seeds and the air in the interstitial spaces among the seeds.
It has been reached when the net movement of moisture from air to
seed, or from seed to air, is zero.
Equilibrium moisture content of seeds is usually determined by the
static method. By this method seeds are placed in containers to hold the
seed sample and acids or dissolved chemicals to provide previously
calculated relative humidities. The containers must be absolutely sealed
from the outside atmosphere and preferably allow entry to measure the
relative humidity of the air. Laboratory desiccators are commonly used
for this purpose. The seeds remain in the container with a known air
relative humidity until equilibrium is reached, after which the seed
moisture content is determined by a reliable method (Hlynka and
Robinson, 1954).
Since the static method may require from a few days to 2-3 months
for seeds to reach equilibrium moisture content with the atmosphere,
another method that reduces this time interval has been developed. By
the dynamic method the air is kept in motion and circulated around the
sample. In one modification of the procedure, the air is passed through
absorption towers, which contain solutions of salts and acids calculated
to control the relative humidity surrounding the seeds. In addition,
Brewer and Butt (1950) found an electric hygrometer to be a useful
instrument for determining equilibrium moisture values within 24 to 72
hours. This is basically a desorption method. Seeds of known moisture
contents are placed in sealed containers with dry air for a specific time.
Thus, moisture moves from the seed to the air in the container. The
relative humidity of the air is determined with the electric hygrometer,
a hair hygrometer, or other means. According to the authors, their
results compared favorably with those obtained by the static method
previously described. The electric hygrometer has the disadvantage of
requiring seed of a wide range of moisture contents, some below the
ambient relative humidities.
Haynes (1961) described a vapor pressure method for determining
seed hygroscopicity. He plotted hygroscopic curves for several seed
species from bis data. His curves appear similar to those plotted from
data obtained by the static method. The method has several disadvan-
tages and limitations. For these reasons and because it has not been
widely accepted, it is not described here.
Hygroscopic equilibrium measurements are usually reported as hav-
ing been made at 25° C, although exceptions are common. The percent-
age of moisture the seeds attain at a given relative humidity, usually at
25°, is known as the moisture content equilibrium.
The equilibrium moisture content of several crop species at different
relative humidities is given in tables 5-9. In tables 7-8, only limited data
were available for forage seeds and most were developed for special
conditions. Dexter (1957) used relatively high humidities to study the
effects of mold formation on equilibrium moisture values for seeds of
several forage species, whereas Harrington (1968) used low humidities
for applications relating to safe sealed storage for at least 3 years.
Unfortunately discrepancies exist between Harrington's (1968) data at
45-percent relative humidity and Dexter's (1957) data at 55-percent
relative humidity and to some extent at 65-percent relative humidity.
Nevertheless these are the best available tables for forage seeds.
Although the data should not be regarded as exact equilibrium moisture
values, especially for research purposes, they can be used safely as
general guides.
TABLE 5.—Equilibrium moisture content of vegetable seeds at various

relative humidities and approximately 25° C- wet basis1
TABLE 6.—Adsorbed equilibrium moisture content of grain seeds at

various relative humidities and approximately 25° C—wet basis'
The sigmoid chape of a typical curve relating relative humidity to

seed moisture content is shown in figure 3. The curve was derived from
average moisture contents and relative humidities of seeds of 10 vege-
table crop species by Nakamura (1958). Because the seeds of each
species have their own moisture-absorbing and moisture-holding po-
tentialities, these curves can be expected to vary to some extent.
Tables 5-9 take into consideration the kinds of seeds and relative
humidity, but they have not been calculated to correct for temperature.
Most research relating to temperature effects has shown that as tem-
perature is increased when seeds are held at a constant relative humid-
ity, the seed moisture content is decreased. Among the researchers who
showed the relationship of temperature to seed moisture equilibrium
were Gane (1941), Gay (1946), Henderson (1952), Thompson and Shedd
(1954), Hogan and Karon (1955), and Hubbard et al. (1957).
Toole et al. (1948) published the seed moisture content of 15 vegetable
species stored at 3 relative humidities and 3 temperatures. The results
obtained at about 80-percent relative humidity and 11.1°, 21.7°, and
26.7° C permit a comparison of temperature effects. In every instance
TABLE 7 .—Equilibrium moisture content of seeds of grasses and

small-seeded legumes at relative humidities below 50 percent and
approximately 25° C—wet basis1
the seed moisture content at 21.7° and at 26.7° was lower than at 11.1°.
However, without exception the moisture content at 26.7° was higher
than at 21.7°. The authors indicated that they had no explanation for
TABLE 8.—Equilibrium moisture content of seeds of grasses and

small-seeded legumes at relative humidities aboye 50 percent and
approximately 23° C—wet basis1
this disparity except possibly "the humidity readings in these chambers

failed to represent the true humidity surrounding the seeds." The
authors pointed out that "the study was not planned to determine the
effect of temperature on moisture content of seed, because the recorded
mean air humidities may not accurately indicate the true conditions
surrounding the seeds." Regardless of the accuracy of the individual
values recorded by Toole et al., the data for 15 vegetable species
strongly indicate that as temperature is increased, equilibrium moisture
content is decreased.
By reducing the results reported by some of these investigators to a
common denominator, the following average values were calculated:
Average decrease in equilibrium moisture content for each 10° C
TABLE 9. —Equilibrium moisture content of seeds of several crops at various relative humidi-
ties and approximately 12° to 25° C1
increase in temperature--wheat 0.65 (Gane, 1941), 0.43 (Gay, 1946),

wheat and corn 0.72 (Thompson and Shedd, 1954), 0.73 (Hubbard et al.,
1957), blue lupine 0.90, corn 1.4, crimson clover 0.62, and sorghum 0.3
(Haynes, 1961). Using the data of Toole et al. (1948) for 15 species of
vegetable seeds stored at approximately 80-percent relative humidity,
the average difference per 10° calculated for storage temperatures of
11.1° and 26.7° amounted to 0.21-percent moisture content. For most of
these croes these relative humidity values do not change greatly as the
temperature is increased or decreased. Although the moisture content
of the seeds increases with an increase in relative humidity, these
values are not greatly affected by temperature.
The slight effect of temperature on seed moisture content is illus-
trated in figure 4 for sorghum and wheat. Although the curves for these
two species are different, the lines for the different temperatures are
parallel, indicating a continuing relationship as humidity is increased.
These relationships do not hold true in all instances. Curves for corn and
rescuegrass seed (fig. 4) show how the curves can deviate as humidity is
increased. For corn, the equilibrium moisture content decreases 0.96
percent for each 10° C increase at 30-percent relative humidity, but at
70-percent relative humidity it decreases 2.3 percent for each 10°
increase. At a relative humidity of about 56 percent the seed moisture
content of rescuegrass seed is the same at -1° and at 49°. As the
FIGURE 4.—Equilibrium relative humidity curves for sorghum, wheat, corn, and rescue-
grass (top to bottom). (Courtesy of Haynes, 1961.)
FIGURE4.—Equilibrium relative humidity curves for sorghum, wheat, corn, and rescue
grass (top to bottom) (Courtesy of Haynes, 1961.)—Continued .
relative humidity is deereased below about 54 percent, the seeds at 49°

hold less water than seeds at -1°. The difference at 25-percent relative
humidity is approximately 0.2-percent moisture for each 10°. On the
other hand, as the relative humidity is increased aboye 56 percent, the
seeds hold more water at 49° than at -1°. The difference at 80-percent
relative humidity amounts to approximately 0.44-percent moisture for
each 10° change.
Rate of Moisture Absorption and Movement

Studies on moisture absorption and movement ir grains and seeds fall
into three categories: (1) Laboratory studies on small samples, (2)
studies on bulk samples of a few pounds to several toas, and (3) studies
on moisture movement of mixed grain with different moisture contente.
For information on moisture movement in grain, see Fisher and Jones
(1939).
Much of the research on equilibrium moisture in small seed samples
has been concerned with ascertaining the time required for the seed to
reach equilibrium. This is regulated by the time required for moisture
(1) to penetrate the seedcoat and (2) to transfer within the seed, based
on the assumption that the relative humidity surrounding the seeds is
uniformly distributed. Since seeds of different species differ as to
seedcoat permeability to moisture, as 411 as constituents of the en-
dosperm and embryo, it is only logical that the time required to reach
equilibrium moisture will vary with the species.
Dillman (1930) found that dry flaxseeds absorbed hygroseopic mois-
ture more rapidly than wheat seeds, which absorbed moisture more
rapidly than alfalfa seeds. Likewise, Pixton and Warburton (1968)
reported similar results with the soft wheat cultivar Cappelle, which
reached 90-percent equilibrium moisture content in less time than did
the hard wheat cultivar Manitoba.
Temperature also affects the rate of water absorption. Dillman (1930)
found the absorption by dry corra, flax, and wheat seed to be twice as
rapid at 30° C as at 10° but no change from 30° to 40°. Whitehead and
Gastier (1946-47) determined that at 20- to 80-percent, relative humid-
ity, sorghum and wheat seeds reached equilibrium moisture content
within 15 days at 20°, whereas nearly 70 days were required at 1°.
The difference between the vapor pressure of the seeds and that of
the surrounding atmosphere is an extremely important factor in deter-
mining the rate of moisture movement. For example, if rice of 13-per-
cent moisture content is placed in an atmosphere of 95-percent relative
humidity, it will increase in moisture content faster than at 70-percent
relative humidity. Likewise, the rice will Tose moisture more rapidly if
placed at 20- than at 50-percent relative humidity.
Pixton and Warburton (1968) measured the rate of hygroscopic
moisture absorption of two cultivara of wheat at relative humidities

from 35 to 90 percent. They found that the time required to reach
equilibrium moisture was neither a gradual progression from 35- to
90-percent relative humidity (absorption) nor a gradual degression from
90 to 35 percent (desorption). Rather, equilibrium was reached in the
following order of relative humidity percentages, with the first number
being the fastest rate:
Absorption-35, 45, 80-90, 60-80
Desorption-85-90, 60-80, 35, 45
Moisture uptake is rather rapid for the first 2 to 3 days for many kinds
of seeds, after which it decreases. According to Babbitt (1949), wheat,
whole or with pericarp and seedcoat removed, reached near equilibrium
moisture content within 40 hours and the increase thereafter was slow.
Results reported by Pixton and Warburton (1968) for wheat more or
less confirmed Babbitt's report. They found that 90 percent of equilib-
rium moisture content was reached within 5-14 days by absorption and
2-9 days by desorption. On the other hand, Breese (1955) indicated that
at a relative humidity from 10 to 90 percent, rough rice required at least
60 days to reach equilibrium moisture. Small samples of chaffy grass
seeds appeared to come to equilibrium moisture content within 3-5 days
over a wide range of relative humidity (Dexter, 1957). Harrington and
Aguirre (1963) dried seeds of six vegetable species to near 0-percent
moisture content and placed them in a saturated atmosphere. After 64
hours they had regained moisture to reach the following moisture
percentages (wet weight basis): Onion 9.4, carrot 9.4, cabbage 8.3,
tomato 8.1, lettuce 8.0, and muskmelon 7.5.
Caution should be exercised in placing much confidence in data
obtained at high relative humidity, especially when observed over long
periods of time. Under these conditions, the changes in weight of
samples may be affected not only by changes in water content of the
seeds but also by the growth of micro-organisms and by respiration.
Movement of moisture finto or out of bulk grain or seed is very slow.
The time for moisture to enter and be distributed within a seed is only a
small fraction of the time required for it to move through a mass of
seed. Instead of the moisture moving from seed to seed or from grain to
grain, it penetrates the mass through the interstitial airspaces among
the seeds. The movement has been measured. Equilibrium was reached
in wheat at a depth 1 inch below the surface after 400 hours (Babbitt,
1949). Seeds of the six species of vegetables used by Harrington and
Aguirre (1963), previously discussed, had an average moisture content
of 8.5 at the surface of the bulk after 64 hours but only 0.7 percent 3
inches below the surface. Frey (1960) found seeds of wheat and birds-
foot trefoil at the center of 100-kg bags to contain 2-4 percent more
moisture than seeds taken from the surface just inside the walls of the
jute bags. The seeds had been stored at five temperature-relative

humidity conditions prior to sampling.
These data show how slowly seeds and grain at the center of large
bulks come to equilibrium moisture content. Were this not so, large
bulks of grain within or outside of enclosures could not be stored safely.
Although the relative humidity surrounding a bin of grain may change
gradually or abruptiy, the outer layers of the grain insulate the inner
layers to a degree. Each layer forms a temporary barrier, which
mediates the change of relative humidity in the next layer. With
continuing high humidity, the barrier is pushed farther toward the
center of the bulk until equilibrium is reached in the entire lot.
A difference in temperature between two areas or zones of a storage
bin or silo can cause seed or grain to deteriorate. When warm air moves
to a colder zone and becomes chilled to the dewpoint, the moisture in the
air is deposited on the seeds as liquid. This situation can exist when
seeds lie in a zone between a cold outside wall of a storage structure and
the interior core where the temperature is higher. Christensen (1970)
demonstrated this principie by using grain sorghum with a moisture
content of 14.3 percent. A temperature difference of 12°-14° over a
distance of 6-8 incites resulted in rapid transfer of moisture from the
warmer to the cooler part of the grain. The grain where moisture
accumulated became heavily infested with storage fungi and decayed.
Equilibrium moisture content was attained within 12 days.
Effect of Extreme Desiccation on Viability and Vigor
Since increased knowledge and improved technology have promoted
the drying of seeds to very low moisture levels for storage, shipment, or
both in sealed containers, it is important to know whether the drying
process causes losses in viability and vigor. There are essentially no
reports of seed damage caused by drying to approximately 6-percent
moisture content, but several workers have reported damage when
seeds were dried to 5 percent or lower. Toole and Toole (1946) reponed
an increase in abnormalities of soybean seedlings grown from seeds
stored for 3 months at a moisture content slightly aboye 5 percent.
Abnormalities consisted primarily of deep tracks across the cotyledons
that interfered with using the stored food reserves.
Roberts (1959) found that timothy seed retained its viability better in
storage as its moisture content was decreased to about 7 percent, but
storage at 5 percent decreased viability. Ching et al. (1959) found a
similar trend with perennial ryegrass seeds. At 6-percent moisture
content they did not retain their viability as well as at 8.3 percent.
Apparently caution must be exercised in storing seeds of some
cultivars of cotton, sorghum, and various other crop species at very low
seed moisture contents. Phillis and Mason (1945) reported that as the
relative humidity of the storage atmosphere was decreased below 40

percent, germination capacity decreased after storage for 5 years at
24°-27° C. Seeds stored at 40-percent relative humidity germinated 75
percent, whereas seeds at 30-, 20-, and 0-percent relative humidity.
germinated 70, 65, and 22 percent, respectively. Seed moisture content
ranged from 6.9, to 5.2, to near O at 40-, 20-, and 0-percent relative
humidity, respectively. Phillis and Mason (1945) found that by placing
the low moisture content seeds at a slightly increased relative humidity,
thus gradually increasing seed moisture content, they could overcome
the deleterious effects of desiccation.
Nutile (1964a, 1964b) found that sorghum seeds that had been dried
to 3.0- to 3.5-percent moisture content showed delayed germination and
produced seedlings with damaged radicles, which subsequently devel-
oped many secondary roots. However, he was able to overcome the
apparent injury by placing the dry seeds at 55-percent relative humidity
until their moisture content increased to 11 percent. Such precondi-
tioned seeds germinated normally.
McCollum (1953), Pollock et al. (1969), and Pollock and Manalo (1970)
noted that cotyledon cracking in garden bean seeds can be caused by
stresses during imbibition and germination as well as by mechanical means.
Very dry bean seeds that developed high percentages of co-
tyledon cracking with rapid imbibition developed less cracking when
allowed to gradually absorb moisture to about 12 percent before im-
bibition.
Much of the damage previously considered as desiccation and me-
chanical damage appears really to be stress damage. If so, seeds can be
dried to very low moisture levels for safe sealed storage, but sensitive
kinds may require special handling prior to planting. Seeds dried to a
very low moisture content for sealed storage apparently must be
allowed to equilibrate with normal atmospheric conditions before
planting except in very arid aneas, where special slow imbibition tech-
niques will have to be employed.
Seeds of some species that can be dried safely to 2- to 3-percent
moisture content are injured when dried to about 1 percent or lower. At
such low moisture content the first indication of injury is reduced rete of
germination, followed in storage experimente by decreased germina-
tion. With severe drying, the symptoms of injury may appear immedi-
ately after drying. As the critical point of damage is approached but not
reached, the symptoms will be apparent after storage. These general-
izations are based on data by Harrington and Cracker (1918). Seeds of
Kentucky bluegrass reduced to 1.5-percent moisture content suffered
no loss either in germination capacity or in vigor. With 0.2-percent
moisture content no loss in germination was evident, but vigor was
reduced considerably, whereas at 0.1-percent moisture content germi-
nation capacity had dropped by 5 percent and vigor was seriously

reduced. Further reduction of the seed moisture content reduced the
germination percentage by one-half and all seedlings were very weak.
This trend has been found for seeds of several crop species but not
Effect of extreme desiccation on seed viability and vigor is shown in
table 10.
Went and Munz (1949) subjected seeds of 113 plant species native to
California to extreme desiccation over phosphorus anhydride (P 205) and
sealed them under vacuum in small gima tubes. No tube was opened
until the appointed time for testing. After 1 year of storage, five species
had lost three-fourths of their germination capacity and some others
had declined to a less extent. It is not known whether this loss was due
to desiccation or to some other factor.
Based on current information, seeds of most crop species can be dried
to 2- to 3-percent m ,isture content without significant injury provided
injury is not caused by another factor associated with the drying
process, such as high temperature. However, drying below 3- to 4-per-
cent moisture content is not recommended for storage of seeds in
commerce, because extremely dry seeds may be damaged by too rapid
rehydration when planted.
Symptoms of desiccation injury may not be apparent until after
storage. They include cracked cotyledons, damage to food transport
system in the embryo, stunted radicle with unusually heavy develop-
ment of secondary roots, stubby primary root and shoot, protrusion of
radicle without further development, and decreased germination com-
pared with nondamaged seeds. The last symptom is usually an excellent
indication of reduced vigor.
Another precaution to observe when drying seed is the possibility of
inducing dormancy. Nutile and Woodstock ( 1967) induced 10- to 25-
percent "low-temperature" dormancy in seeds of six sorghum cultivars
by drying. When placed to germinate at 15° C, dormancy was apparent,
but at 25° the blocks to germination did not exist. Species, and possibly
some cultivars, for which experimental data are not available might well
be treated as suspect.
Interrelationship of Temperature, Seed Moisture

Content, and Storage Life
To some degree both temperature and seed moisture content can be
controlled for practical storage. The nature and degree of control will
depend on the economice applicable to specific situations. These deci-
sions may be affected by local climate and the quantity of seed to be
stored as well as by the length of storage.
Available data on storage of hempseeds demonstrate the effects of
TABLE 10.—Effect of extreme desiccation on seed viability and vigor of several crop species
TABLE 10. —Effect of extreme desiccation on seed viability and vigor of several crop species—Con.
temperature and moisture on storage life. In 1950, seed lot 2 was grown
in Kentucky and seed lot 4 in Maryland, and in 1951 they were stored at
Beltsville, Md., until 1960-61, when they were transferred to the
National Seed Storage Laboratory, Fort Collins, Colo. The tests over
the first 51/2 years were malle at Beltsville by Toole et al. (1960) and
those from 9 to 12 years of storage by Clark et al. (1963). Lot 2
represented healthy seed of strong vigor, which had a germination
capacity of percent when stored. The viability of lot 4 when stored
was approximately 88 percent. There was no significant losa in germi-
nation of lot 2 stored in sealed containers at 6.2-percent moisture,
Unfortunately the data are not available for seed stored at 6.2-percent
moisture and 21° C beyond 51/2 years.
Table 11 shows the effect of seed moisture content, temperature, and
length of storage on seed germination. Seed lot 2 stored at 6.2-percent
moisture content and 10° C germinated 96 percent after 12 years,
whereas seed stored at 9.5-percent moisture content and 10° lost its
viability completely during the same period. Seed stored at 9.5-percent
moisture content and 10° required 11 years to reach 5-percent germi-
nation, but seed stored at the same moisture content and 21° required
only 2 years to reach this germination level. There is greater variation
among germination values for lot 4. This is common in seed lots of
declining viability or of low vigor. Otherwise the trends for moisture
and temperature effects are comparable for the two seed lots.
The effect on an oilseed crop, soybean, of a combination of high
temperature (30° C) and high moisture content (18.1 percent for Mam-
moth Yellow and 17.9 percent for Otootan) is given in table 12. Only 14
percent of the Mammoth Yellow seeds and none of the Otootan seeds
were viable after 1 month of storage. The high moisture content of both
cultivars stored at -10° caused little or no deterioration until after 72
months of storage. These data show that the viability of soybean is
maintained longer when stored at subfreezing temperatures than at 2°.
No seeds of either cultivar with 9.4- and 8.1-percent moisture content
decreased in viability when stored at -10°, 2°, or 10° for 10 years (cols.
2-4).
The results of an experiment with nine species of vegetable seeds and
peanut for storage periods of 110 and 250 days are given in table 13.
This table differs from tables 11 and 12 in that it is based primarily on
vegetable seeds, evaluates storage conditions by crop species and by
their averages, and concentrates on intermediate temperatures. Al-
though the moisture contents at which the seeds were stored are not
shown in table 13, they can be estimated for the different crop species at
each humidity level by referring to tables 5 (veg.) and 9 (peanuts).
The average results for both storage periods show that as storage
temperature or storage relative humidity is inereased the germination
TABLE 11.—Germination of 2 lots of hempseed stored in sealed con-

tainers at 8 temperature-moisture conditions for 12 years1
of the seeds is reduced. After 110 days the averages of seeds stored at
10° C and 81-percent relative humidity differed by only 0.6 percent from
those at 26.7° and 44-percent relative humidity. However, after 250
days the averages differed by 4.1 percent. The data for 44- and
78-percent relative humidity at 26.7° show that seeds of tomato retained
good germination longest and seeds of onion and peanut the shortest
time.
Barton (1939) in experiments with aster seeds over 3 years reported
on seed deterioration under several storage conditions. One of the most
important resulta in table 14 is the consistently high germination of the
seeds stored under controlled conditions of low temperature and low to
moderate seed moisture content compared to the relatively rapid de-
terioration of seeds stored at ambient temperature and the same

moisture content.
Vacuum and Gas Storage

For many years research has been conducted on the effects of a
partial vacuum and such gases as carbon dioxide, oxygen, and nitrogen
on the longevity of various kinds of seeds. The reported results from
these studies are variable and in some instances appear contradictory.
This confusion undoubtedly results from the widely divergent test
methods employed by the various researchers. Because of the lack of
complete information on the test procedures used in numerous studies,
direct comparisons cannot be made between and among the various data
found in the literature. Likewise sealed storage cannot be directly
compared with open storage, because in sealed containers oxygen
concentration in the atmosphere decreases and the carbon dioxide
concentration increases with time (Harrington, 1963), whereas in open
storage the composition of the atmosphere remains constant. Because it
is not feasible to continually adjust the composition of the atmosphere in
sealed containers, most studies have not included gas analyses.
Some workers paid no attention to either seed moisture or storage
temperature, whereas others attempted to control one or the other.
Several used air-dry seeds and room temperature, both of which
provide a minimum of information about the conditions actually used.
Regardless of the kind of seed, the air-dry moisture content in one
geographic area is not necessarily the same as that in another. Likewise
room temperature is not consistent from place to place nor from month
to month, or even from day to day in most localities. To accurately
assess the protective value of either a partial vacuum or a gas, all
environmental conditions have to be considered. Reports on a few kinds
of seeds will illustrate the variable results in the literature.
Barley
The higher the oxygen content of the storage environment, the
shorter the viability period for barley seeds (Roberts and Abdalla,
1968). The deleterious effects of oxygen are produced at relatively low
oxygen concentrations and are most pronounced at the higher seed
moisture levels.
Corn
Since the seedsman wants to store his seeds at the highest practical
moisture leve) and temperature, Goodsell et al. (1955) stored seed of
two corra hybrids at approximately 8-, 10-, 12- and 14-percent moisture
sealed in air, carbon dioxide, and nitrogen at approximately —18°, 4°,
16°, and 29° C. Generally the average pooled germination of the two
TABLE 12.—Seed germination of 2 soybean cultivare stored at 2 moisture contents and 5 temperatures for 10 years1
1Data from Toole and Toole(1946).
2Mammoth Yellow 9.4 and Otootan 8.1 percent.
3 Mammoth Yellow 13.9 and Otootan 13.4 percent.
4 Mammoth Yellow 18.1 and Otootan 17.9 percent.
5 Mammoth Yellow 97- and Otootan 93-percent germination before storage.
TABLE 13. —Seed germination of 10 crops after 110 and 250 days of storage at 6 temperature-relative
humidity conditionsl
See footnotes at end of table.

TABLE 13. —Seed germination of 10 crops after 110 and 250 days of storage at 6 temperature-relative
humidity conditionsl —Continued
TABLE 14.—Germination of aster seeds after storage at 12 tempera-

ture-moisture conditions over 36 months 1
hybrids in the three storage gases was still 95 percent or better after 5
years for the seeds at 8-, 10-, and 12-percent moisture at –18° and 4°
and those containing 3- and 10-percent moisture at 16°. Seeds containing
12- or 14-percent moisture at 16° and those with 8- and 10-percent
moisture at 29° deteriorated rapidly after 1 year. Corn seeds containing
12- to 14-percent moisture were practically all dead after one-half to 1
year, regardless of the surrounding gas.
Sayre (1940) stored corn seeds with 18-percent moisture oxygen,
carbon dioxide, and nitrogen at 30° . The seeds in oxygen died within 3
years and the germination of the seeds in carbon dioxide and nitrogen
dropped noticeably. At low temperatures corn seeds with 18-percent
moisture sealed in carbon dioxide and nitrogen had good germination
for 5 years.
Struve8 dried corn seeds to near 0-percent moisture, sealed them in
oxygen and in nitrogen, and stored them at –30° to 50° C. Seeds at –30°
remained unchanged through 31 months, those at 50° in nitrogen

showed a progressive reduction in vigor, and seeds in oxygen died
within 7 months. He also stored corn seeds with 1-percent moisture
content i,i atmospheres containing 0-, 20-, 60-, and 100-percent oxygen
at 0°, 5°, 10°, 15°, 25°, 35°, and 50°. He concluded that oxygen concen-
tration may become an important factor in the deterioration of corn
seeds in sealed storage.
Flower Seeds
Primula sinensis sealed in carbon dioxide declined only 30 percent in
viability over a 7-year period, whereas unsealed seeds lost all viability
(Lewis. 1953). Seeds of Salvia splendens deteriorated seriously when
sealed under a vacuum (Chopinet, 1952). Aster seeds kept equally well
when sealed in air or a partial vacuum (Barton, 1939). Sealing under a
partial vacuum had no advantage over sealing in air for maintaining the
viability of verbena seeds (Barton, 1939).
Barton (1960a) stored seeds of Lobelia cardinalis with 6.9- and
4.7-percent moisture ín sealed glass vials in air, oxygen, carbon dioxide,
nitrogen, and vacuum and in open containers for up to 25 years at
laboratory temperature, 5°, and —5° C. In all cases, viability was lost
more rapidly at laboratory temperature than at 5° and —5° and at
6.9-percent than at 4.7-percent moisture. At laboratory temperature,
6.9-percent moisture seeds in air, open or sealed, lost viability rather
rapidly. Seeds sealed in oxygen lost viability even more rapidly,
whereas carbon dioxide, nitrogen, and a partial vacuum extended the
life of the seeds for 6 to 8 years. When seed moisture was reduced to 4.7
percent, seed longevity was increased at laboratory temperature. Car-
bon dioxide, nitrogen, and vacuum were superior to air and oxygen,
with oxygen causing the most rapid deterioration, There were no
significant differences in the response of seeds to atmospheres of carbon
dioxide, nitrogen, or under vacuum at room temperature or to air,
oxygen, carbon dioxide, nitrogen, or vacuum at 5° and —5° except those
resulting from increased time in storage.
Grasses
The effect of nitrogen was negligible on the longevity of Chewings
fescue (Gane, 1948a) and meadow fescue (Evans et al., 1958). There
was no advantage in using either nitrogen or a partial vacuum rather
than air for sealed storage of seeds of Kentucky bluegrass and creeping
red fescue (Isely and Bass, 1960). In fact, when the seeds were
subjected to an unfavorable temperature, loss of viability was more
rapid for seeds packaged with nitrogen or under vacuum than with air.
Legumes
Seeds of both red and white clover stored under vacuum and in
nitrogen were shorter lived than those stored unsealed (Davies, 1956).
Red clover seeds containing 10.3-percent moisture when sealed with
carbon dioxide lost all viability in 23 years, but when calcium ehloride
was used with the carbon dioxide, only about one-third of the bala'
viability was lost (Evans, 1957a).
No atmosphere tested, including air, vacuum, carbon dioxide, nitro-
gen, helium, and argon, was consistently or significantly better than all
others for 2 years of sealed storage of crimson clover seeds (Basa et al.,
1963a).
Oilseeds
Soybeans in open storage for nearly 6 years lost viability (Guillaumin,
1928), whereas seeds sealed in an atmosphere free of oxygen germi-
nated 92 percent and those under a vacuum had 100-percent viability.
Low moisture (7 percent) cottonseeds retained their initial viability
when sealed in air, oxygen, carbon dioxide, or nitrogen and stored at 21°
and 32° C (Simpson, 1953). Seeds with 13-percent moisture dropped to
one-half to two-thirds of the original germination under all storage
conditions. The loss of germination with oxygen was no greater than
with carbon dioxide or nitrogen; however, the loss in air was greater
than in the pure gases.
Russ et al. (1963b) found that air, vacuum, carbon dioxide, nitrogen,
helium, or argon was neither consistently nor significantly better than
the others for sealed storage of safflower and sesame seeds for 2 years.
Rice
Much of the literature pertains to the storage of rice seeds for both
food and seed. In areas where rice is grown, seed moisture content
tends to remain high even when the seeds are air-dry. Deterioration of
high moisture (20.8 percent) seeds at 30° C can be delayed for a few
weeks by sealing them in an atmosphere of carbon dioxide mixed with
1,000 p/m of ethylene oxide (Kaloyereas, 1955).
Kondo and Okamura (1927, 1929, 1930 1934) and Kondo et al. (1929)
found that both rough and hulled rice can be stored sealed with carbon
dioxide or air for up to 4 years with little loss of viability provided the
seed moisture content is less than 13 percent. They reported that
carbon dioxide had a slight advantage over air. Rice dried to 5-percent,
moistum and sealed in an atmosphere of nitrogen germinated 99 per-
cent alter 8 ycars, but with 13-percent moisture all viability was lost
(Sampietro, 1931). Seeds with either 5- or 13-percent moisture lost all
viability when sealed in carbon dioxide, air, or under a partial vacuum.
Sorghum
Sorghum seeds during the second year of storage retained signifi-
cantly higher germination when sealed under a partial vacuum than
when sealed in air, carbon dioxide, nitrogen, argon, or helium (Basa et
al., 1963a).
Vegetable Seeds
For pea seeds the period of safe storage decreased as the oxygen
concentration in the storage atmosphere increased from O to 21 percent
(Roberts and Abdalla, 1968). The deleterious effects of oxygen were
more pronounced at the higher seed moisture contenta. There was no
advantage in using sealed storage under nitrogen or a partial vacuum
rather than air for onion seeds (Gane, 1948b; Isely and Basa, 1960).
However, seeds sealed in carbon dioxide retained their viability better
than did similar seeds sealed in air (Lewis, 1953; Harrison and McLeish,
1954; Harrison, 1956). Vacuum, carbon dioxide, nitrogen, helium, and
argon storage had no advantage over sealed-in-air storage for lettuce
seeds during 2 years (Basa et al., 1962). Although lettuce seeds sealed
in carbon dioxide retained their viability better at room temperature
than did similar seeds sealed in air, storage in carbon dioxide revealed
differences in longevity between cultivara (Harrison and McLeish,
1954; Harrison, 1956).
Cabbage seeds stored equally well when sealed in air, nitrogen, or a
partial vacuum (Isely and Bass, 1960). Parsnip seeds retained their
viability better when sealed in carbon dioxide than in air (Lewis, 1953;
Harrison, 1956). Dandelion seeds retained their viability about equally
well when sealed ir partial vacuum or in air, except seeds containing
7.9-percent moisture seemed to deteriorate more rapidly in a partial
vacuum (Barton, 1939).
Wheat
Wheat, like rice, is an important food crop, which has received much
attention because the condition of storage greatly affects its milling and
processing qualities as well as its seed germination. To improve the
storage of wheat seeds for both planting and food purposes, studies
have been made on the effects of nitrogen and carbon dioxide on seed
viability. Loss of viability of high moisture wheat seeds can be delayed
for several days by sealing under either 50 or 75 percent of carbon
dioxide (Peterson et al., 1956) or nitrogen (Glass et al., 1959). However,
once deterioration starts, it proceeds rapidly. It can be delayed for
several additional weeks by combining nitrogen storage with a lower
temperature (20 0 C); however, even this combination is unsatisfactory
for extended storage.
PRINCIPLES AND PRACTIES OF SEED STORAGE
Confirmation Studies at the National Seed Storage Laboratory

Obviously some kinds of seeds under certain circumstances are ben-
efited by vacuum or gas storage. The question then is "Under what
conditions is vacuum or gas storage practical and desirable?" To more
fully understand the interrelationship involved, a comprehensive study
was undertaken at the National Seed Storage Laboratory, Fort Collins,
Colo. One lot each of the following kinds of seeds was adjusted to 4-, 7-,
and 10-percent moisture and sealed in air, under vacuum, and in carbon
dioxide, nitrogen, helium, and argon and stored at –12°, –1°, 10°, 21°,
and 32° C: 'Dixie' crimson clover, 'Great Lakes' lettuce, 'Pacific No. 1'
safflower, `Margo' sesame, and `RS 610' hybrid sorghum. No gas analy-
ses were made after storage. Germination tests were made at the time
of storage and at intervals thereafter. Because commercially available
equipment was very expensive, Bass and James ( 1961) developed a
simple, inexpensive device for vacuum and gas sealing of tin cans. Terne
complete sealer (fig. 5) consists of a vacuum chamber, soldering gun,
three-way valve, vacuum gage, several lengths of air hose, hose clamps,
a brass tee fitting, and a source for both vacuum and gas.
Germination tests were made in electronically controlled water-cur-
tain germinators operated at 20° C for crimson clover and lettuce and a
20° to 30° night-to-day alternation for safflower, sesame, and sorghum.
Each test consisted of two 100-seed replicates planted according to
official procedures (Association of Official Seed Analysts, 1954) except
safflower, for which official procedures had not been developed at that
time. Safflower was planted the same as sorghum. In all tests only
normal seedlings—those capable of producing plants—were recorded.
For crimson clover, which has hard seeds, the percentage of hard seeds
was added to the percentage of normal seedlings to give total germina-
tion.
The data (table 15) show that regardless of the kind of seed, no
storage atmosphere consistently gave the highest germination percent-
age at all temperatures for seeds of all moisture levels tested. The data
also show that there are distinct differences between kinds of seed in
their response to temperature and seed moisture content.
Of the five kinds of seeds, sorghum showed the least sensitivity to the
interaction between seed moisture content, storage temperature, and
storage atmosphere (table 15). Only the 10-percent moisture seed
showed really drastic germination reductions when stored at 32° C. At
that temperature seeds in all atmospheres except those under a partial
vacuum (3 percent) and in argon (5 percent) were completely dead. At
21°, seeds with 10-percent moisture in a partial vacuum, helium, and
argon did not show a significant reduction in germination, but seeds in
all other atmospheres did. Significant reductions in germination were
recorded for 4-percent moisture seeds under vacuum at 21' and in air,
nitrogen, and helium at 32°. Significant reductions in germination oc-
curred for 7-percent moisture seeds in all atmospheres at 32°. No
significant reductions were recorded for seeds of any moisture content
in any atmosphere at 10°, —1°, and —12°.
Crimson clover seeds (table 15) stored in paper envelopes (check)
showed a significant decrease in germination after 8 years of storage at
each temperature for each initial moisture level except 4- and 10-per-
cent moisture seeds held at —12° C. Except for 7- and 10-percent
moisture seeds at 10°, 21° and 32°, only an occasional sample of sealed
seeds showed a significant decline in germination. Such declines were
not consistent for storage temperature, seed moisture, or surrounding
atmosphere. A significant decline in germina ion of sealed seeds stored
at —12° was recorded for 7-percent moisture seeds in a partial vacuum
and for 10-percent moisture seeds in air. At significant germination
losses occurred with 7-percent moisture seeds held in nitrogen and with
10-percent moisture seeds held in air, vacuum, carbon dioxide, nitro-
gen, and helium. The significant germination declines recorded at 10°
occurred for 7-percent moisture seeds in air, carbon dioxide, and
nitrogen and for 10-percent moisture seeds in air, vacuum, carbon
dioxide, nitrogen, and helium. At 21°, significant decreases occurred for
7-percent moisture seeds in air, carbon dioxide, nitrogen, and helium
and for all 10-percent moisture seeds.
The only significant decrease in germination of 4-percent moisture
seeds held at 32° C was for those in nitrogen. AH 7-percent moisture
seeds showed a significan* decline, with the highest germination main-
tained in an atmosphere of argon. Argon also gave the highest germi-
nation of 4-percent moisture seeds at 32°.
The sealed l0-percent moisture seeds at 32° C showed a significant
decline in germination the first year of storage and germinated 12
TABLE 15. —Germination of 5 kinds of seeds stored with approximately 4-, 7-, and 10-percent moisture content at 5
temperatures in various atmospheres in sealed metal cans; check samples were in paper envelopes
TABLE 15.—Germination of 5 kinds of seeds stored with approximately 4-, 7-, and 10-percent moisture content at 5
temperatures in various atmospheres in sealed metal cans; check samples were in paper envelopes—Con.
TABLE 15. —Germination of 5 kinds of seeds stored with approximately 4-, 7-, and 10-percent moisture content at 5
temperatures in various atmospheres in sealed metal cans; check samples were in paper envelopes—Con.
percent or less at the end of the second year. By the end of the eighth
year, even the few hard seeds initially present were dead.
Maximum germinations were distributed among atmospheres as fol-
lows:
In some tests the same germination was recorded for seeds in two or
more atmospheres. This finding applies to the tabular data for lettuce,
saffiower, sesame, and sorghum (pp. 72-74).
The data for lettuce (table 15) show that seed moisture and storage
temperature had more effect on germination than did the composition of
the surrounding atmosphere. Drying to 4-percent moisture before
sealing made possible the storing of lettuce seeds at as high as 32° C
with significant differences between atmospheres evident only at 32°.
The apparent significant decline to 5-percent germination for 4-percent
moisture seeds in a partial vacuum at 10° resulted from a defective seal
after evacuation. The 1-percent germination for 7-percent moisture
seeds in air at 10° also resulted from a defective seal. In both tests a fine
hole in the solder allowed the seeds to absorb moisture from the
atmosphere until an unsafe level was reached. These data emphasize
the need for extreme care in sealing seed containers. Highest germina-
tion of lettuce seeds in sealed cans occurred as follows:
Safflower seeds must be dried to 4-percent moisture (table 15) for

safe, sealed storage at ordinary temperatures regardless of the sur-
rounding atmosphere. Seeds containing 7-percent moisture cannot be
stored safely above 10° C, and 10-percent moisture seeds cannot be
stored above -1°. All sealed 10-percent moisture safflower seeds
showed a significant loss in germination during 8 years of storage. The
array of highest germinations for safilower seeds sealed in various
atmospheres was as follows:
The data for sesame seeds (table 15) show that this oilseed does not
store well when sealed in any atmosphere at moisture levels above 7
percent unless the temperature is kept at -1° C or lower. For sealed
storage in any atmosphere at higher temperatures seed moisture con-
tent must be reduced to 4 percent. The distribution among atmospheres
of the highest germination percentages for sesame seeds was as follows:
No atmosphere was consistently better than all others for preserving

the germination of sealed sorghum seeds. Therefore it is interesting to
note that among seed moistures and storage temperatures the following
combinations, with atmospheres that gave the highest germinations
after 8 years of sealed storage, were about equally divided:
The distribution of highest germinations for all five kinds of seeds

among the various atmospheres in sealed metal cans is summarized as
follows:
It is obvious from this distribution of highest germinations that no one

atmosphere is decidedly better than the others for safe, sealed storage
of seeds.
The data in table 15 show that regardless of the kind of seeds or the
atmosphere in the sealed container, only adequately dried seeds (
4 percent moisture) retained their germination reasonably well during 8
years of storage at 32° C. Some kinds did not keep well at 21°, and
sesame seeds with 7- and 10-percent moisture lost all viability at 10°
except for 1 or 2 percent when sealed in helium or argon.
It is obvious from the data that seed moisture cortent at the time of
sealing has a far greater effect on seed longevity than does the sur-
rounding atmosphere. For most situations the added expense of using
atmosphere other than air appears unnecessary. However, even for
sealing in air, most kinds of seeds require additional drying before

sealing if the sealed seeds are to be subjected to a temperature above
10° C. In general, the added advantage, if any, of an atmosphere other
than air is not realized except during very long-term storage.
Illumination
Only limited studies have been made on the possible effects of light on
stored seeds. At least four investigators, who have studied the potential
effects of white light on the storage life of seeds, have found no positive
effect of illumination that could not be explained by the reduction in
seed moisture content caused by the light treatment (Tammes, 1900;
Bacchi, 139.5; Wyttenbaeh, 1955; Litynski and Urbaniak, 1958). Har-
rington (1972) indicated that ultraviolet rays from the sun can reduce
longevity of seeds before harvest and pasten deterioration of stored
seeds. Since Harrington (1972) did not indicate the source of his
information, these statements are assumed to be extrapolations from
the effects of ultraviolet radiation on other biological materials.
Barton (1947) claimed that when Cinchona ledgeriana seeds with a
moisture content as high as 9.4 percent were exposed to light during
storage in a laboratory, there was some evidence of harmful effects.
However, the author indicated that the moisture content of the control
samples stored in darkness may have been reduced during storage and
thus provided a more suitable storage environment. Also, seeds stored
in sealed glass tubes and not resealed for future use after opening did
not confirm the initial conclusion.
Perhaps the most comprehensive study of this type was made by
Litynski and Urbaniak (1958). For 4 years they studied light effects on
the storage life of seeds of eight vegetable species and seven field crop
species that had been exposed to white light, orange-yellow light,
blue-violet light, diffused light of reduced intensity, and darkness. In
studying possible benefits of spectral characteristics of the tilters used
and light intensity, they found no consistent benefit. Any apparent
benefit might be explained by the possible drying of the seeds by the
orange-yellow rays as opposed to the lack of drying by the blue-violet
rays. Diffused light was not beneficia) to the retention of seed viability.
Jensen (1941, 1942) claimed that the storage life of cabbage and
cauliflower seeds was increased significantly by illumination simulta-
neously with a mercury-quartz lamp and a heat lamp. Principal irradia-
tion by the mercury-quartz lamp was from 300u to 450u with a max-
imum energy peak at about 325u, whereas the heat lamp emitted
radiation from about 350u to 550u with a peak at 485-550u. In Jensen's
work (1941, 1942)the seeds were exposed to light in the open, either on
a canvas with constant stirring or on a moving belt provided with
ventilation. Although we cannot disagree with the claims that light

affects the storage life of seeds, we believe that the author's findings did
not exclude the possibility of seed dehydration, which resulted from the
treatment procedure responsible for the increased storage life.
Harrington (1972) explained the relationship of a macromolecular
water layer, which insulates a maeromolecule against oxidation and
subsequent deterioration of molecules, cells, and embryonic tissue. The
removal of this macromolecular layer renders the molecule vulnerable
to oxidation by several agents including ultraviolet light. Maes and
Bauwen (1951) studied the effect of infrared rays on preserving wheat
seeds and some oilseeds. They found that infrared raya inhibited the
lipolytic degradation of wheat germ and some oilseeds. The phenome-
non was not caused by a change in the moisture content of the germ, as
the moisture content remained unchanged during radiation and storage.
The published information on the effects of illumination on the
storage life of seeds is controversial. Until these controversias are
resolved, no general recommendation can be made that would apply to
field, vegetable, and fiower seeds.
Respiration and Heating

Respiration has been defined as an oxidative-reduction process oc-
curring in all living cells, whereby chemical action occurs, producing
compounds and releasing energy that are partly used in various life
processes. Respiration is discussed here only as it relates to practical
seed storage. The following phases respiration may be considered:(1)
Depletion of food reserves, (2) formation of intermediate or end prod-
ucts that may affect seed storage, and (3) release of energy, much of
which is in the form of heat. Of these three phases, perhaps the release
of heat has the greatest application to seed storage.
Depletion of Food Reserves
Since respiration is an oxidation procese, there must be a substrate
(the seed) with which the oxygen can combine. Also, respiration is
possible only in the presence of enzymes, some of which are very
specific and some more general in their functions. As respiration pro-
ceeds, more and more of the seed food reserves are used up. There is
some evidente that the carbohydrate reserves in the endosperm of
wheat, on which much research of this nature has been conducted, are
used before the reserves in the embryo. Ordinarily the sugars and
starches are digested before the lipids and proteins. When stored seeds
are protected from insects and vermin and the environment is unfavor-
able for the development of molde and bacteria, the respiration process
continues under some conditions to the state or condition wherein the
seed appears as charcoal, for example, carbonized barley seeds taken
from the Anatolian excavations of Alishar, dating to about 1000 to 3000

B.C. It is doubtful that seeds stored under reasonably favorable condi-
tions of temperature and humidity would lose enough dry weight over 5
to 10 years to significantly weaken them.
Respiration Products That Retard This Process

One product of respiration is carbon dioxide. Most relevant research
has shown that in a closed system the accumulation of carbon dioxide
inhibits respiration (Crocker and Barton, 1953). In bag and bulk storage
the carbon dioxide is usually dissipated and thus has little effect on
respiration. Miiner and Geddes (in Anderson and Alcock, 1954, pp.
152-220) indicated that renewal of oxygen in open storage can occur
with normal changes in barometric pressure. The inhibition of respira-
tion by carbon dioxide in a closed system can be used advantageously by
storing seeds in sealed containers in which the carbon dioxide replaces
the air. Inhibitory concentrations of carbon dioxide in respiring soy-
beans were found to be from 12 to 14 percent. Milner and Geddes (in
Anderson and Alcock, 1954, pp. 152-220) reported that a carbon dioxide
corcentration of 7 percent affected respiration in wheat and at 12
percent inhibition was marked.
Release of Heat in Respiration
The typical textbook equation for respiration is
C 6H 120 6 + 6CO 2 = 6CO 2 + 6H 20 + 673 kg calories.
This equation shows that for oxidatien of 1 mole of a hexose, 6 moles of
oxygen are required; that 6 moles each of carbon dioxide and water
result from this oxidation; and that 673 kg calories of energy are
released. The complete combustion of a gram of glucose yields 3.76
calories. Not all glucose or other sugars proveed to complete combus-
tion. Moreover the amount of heat released per unit of substrate varíes
with the kind of material involved. The important point here is that heat
is evolved from respiration.
Under favorable storage conditions the heat of respiration is of little
or no concern for practical seed storage. However, at higher moisture
levels the heat of respiration can produce much damage to stored seed.
This would probably occur with freshly harvested seed having a high
moisture content (De Witt et al. , 1962), with seed in storage areas not
well protected from the weather, with seed in the holds of ships where
the temperatures are high, or where it is not adequately protected from
the elements.
Measurement of Respiration
Because respiration in the resting seed proceeds very slowly, equip-
ment and methods to measure the amount and rate must be precise.
Methods commonly used to study respiration in seeds include measuring

oxygen absorption and carbon dioxide evolution. Because each method
has weaknesses (see Bartholomew and Loomis, 1967), many re-
searchers measure both oxygen absorption and carbon dioxide evolu-
tion, which also provide data for computing the respiration quotient.
This quotient is the ratio of carbon dioxide produced to the amount of
oxygen consumed, usually written as CO 2/02 . Attempts have been malle
to measure water production in respiration, decrease in dry matter
(substrate), evolution of heat, and loas of energy as a measure of
respiration. These methods have not been as successful for measuring
small bulk samples as measurement of oxygen absorption and carbon
dioxide evolution.
Some Factors Affecting Respiration
The apparent respiration of seed lots and the associated heat may
arise not only from the seeds but also from the fungi and bacteria on the
seeds, especially at seed moisture contents in equilibrium with approx-
imately 75-percent relative humidity and higher. According to Milner
and Geddes (in Anderson and Alcock, 1954, pp. 152-220), critical
moisture percentages that increase respiration are wheat 14.6, barley
14.5, corn 14.2, and flaxseed 11.0. Other examples of equilibrium
moisture content of seeds are in tables 5-9. Since mold spores do not
germinate at relative humidity levels much below 75 percent, the
relative humidity of the storage arca should be considerably leas than 75
percent for sale storage for more than 1 year. Molds grow readily at
humidities above 75 percent. Bacteria are of minor importance in seed
storage as their relative humidity requirement is above 90 percent. If
held at optimum temperature (usually 25° to 30° C), fungus spores may
germinate at a lower relative humidity than at a nonoptimum tempera-
ture.
Respiration increases as temperature is increased until limited or
terminated by inactivation of enzymes or enzyme systems, by increase
in termperature, exhaustion of the substrate or oxygen, and accumula-
tion of carbon dioxide to inhibitory levels. Ching (1961) found that
increased moisture accelerated respiration of perennial ryegrass and
crimson clover seeds more than did increased temperature.
Preservatives for Reducing Respiration

Considerable research has been conducted to find a chemical that will
inhibit the development of micro-organisms on seeds, grain, and oil-
seeds without adversely affecting their quality. Much of this work was
done at the Department's Southern Regional Research Laboratory,
New Orleans, La., by Aitschul et al. (1946) and Lambou et al. (1948).
Preserving seed viability for propagation poses quite a different prob-
lem than controlling the growth of molds on grain and oilseeds to be

used for milling and industrial purposes. Unfortunately no successful
chemical or application has been found to control storage micro-orga-
nisms or respiration without damaging the seed embryo.
Conclusion
The most effective method of keeping the respiration of stored seed
to a minimum is to keep the seed dry. It should be dried to a moisture
content safe for storage and held at a relative humidity that will
maintain a safe moisture content throughout storage. Most of the
research on seed respiration has been on cereal seeds, oilseeds, and tree
seeds. Nevertheless seeds of other plant groups or species have been
the subject of interesting studies. For more information, see Miller
(1938), Crocker and Barton (1953), and Milner and Geddes (in Anderson
and Alcock, 1954).
EFFECTS OF PESTS AND CHEMICALS ON SEED

DETERIORATION IN STORAGE 9
At harvest, seed lots contain much extraneous material, most of

which is removed by cleaning. However, certain fungi, bacteria,
viruses, and insects are not removed, and they cause or hasten seed
deterioration. In addition, chemical treatments to control fungi, insects,
and rodents may affect germination and longevity.
Seedborne virases are rather uncommon. When present they usually
remain dormant in stored seeds and infect the seedling subsequent to
germination. Bacteria and actinomycetes, although capable of damaging
stored seeds, usually remain quiescent because the relatively high
moisture content necessary for their growth seldom occurs in commer-
cial seed storage.
Storage Fungi
Saprophytic and parasitic seedborne fungi include Alternaria , Chae-
tomium, Cladosporium, Fusarium, Helminthosporium, and Rhizopus
spp. They remain dormant during seed storage unless seed moisture
content increases greatly, e.g., to over 14 percent in cereals. However,
these fungi may prevent infection of grass seeds by storage fungi
during the first part of the storage period (Kulik and Justice, 1967).
Certain molds (fungi), not usually present in or on seeds at harvest, can
carry out their lile cycle on stored seeds (fig. 6).
These fungi, termed "storage fungi" by Christensen and Kaufmann
(1965) to differentiate them from the "field fungi," which are present in
or on seeds prior to storage, can destroy stored seeds (table 16), because
they can grow under limited moisture conditions where field fungi and
other micro-organisms cannot grow. In fact, many of the storage fungi
are actually osmophilic and grow best under relatively dry conditions
(table 17). Some can invade seeds with moisture contents in equilibrium
with an ambient relative humidity of 65 percent. Typical moisture
contents are corn 13.8, oats 13.2, rye 14.3, and wheat 14.1 percent
(Kreyger, 1954). (All moisture contents in this section are on a wet
weight basis.)
Storage fungi have been reported to invade and destroy cereal grains
(Christensen and Kaufmann, 1969), cocoa beans (Bunting, 1930), cot-
tonseeds (Arndt, 1946), grass seeds (Kulik and Justice, 1967), locust-
beans and Beach beans (Snow et al., 1944), peas (Fields and King,
1962), peanuts (Diener, 1958), and soybeans (Milner and Geddes, 1946).
They can attack almost any kind of seed under favorable environmental
conditions, since these molds can grow on most organic materials.
Invasion of seeds by storage fungi may result in loss of viability,
increase in free fatty acids, decrease in nonreducing sugars, develop-
ment of musty odors, and discoloration. Deterioration can occur in a few
days when seeds are stored under unfavorable conditions.
Storage fungi are principally Aspergillus and Penicillium spp., which
commonly occur throughout the world. They are usually present in
large numbers in the air and on surfaces in seed storage areas; they may
be on a few seeds in a lot. They will invade and destroy seeds at 4 -45 ° C
and 65- to 100-percent relative humidity. Their activity is largely
determined by the physical condition, vitality, and moisture content of
the seed and the ambient temperature and relative humidity of the
storage area. Consequently, the fungal population reflects the kind and
efficiency of the postharvesting handling, conditioning, and storage
environment of a seed lot.
Much of our knowledge of storage fungi has come from studies of
their activity in stored cereal seeds. Of the many Aspergillus spp., only
a few have been reported to attack stored cereal seeds. A. amstelo-
dami, A. chevalieri, A. repens, A. restrictus, and A. ruber grow in
seeds with a moisture content of 13.2 to 15 percent. The major fungi
found in cereal seeds above 15-percent moisture content are A. can-
didus, A. flavus, A. ochraceus, A. tamarii, and A. versicolor (Chris-
tensen, 1957). Penicillium spp., though not as common as Aspergillus
spp. , are often found in cereal seeds, particularly in lots with a moisture
content above 16 percent and stored at relatively low temperatures
(Christensen and Kaufmann, 1965).
A rapid growth of fungi in stored seeds can produce so-called hot
spots caused by heating. Using seeds with 18-percent moisture content,
Gilman an Barron (1930) found in laboratory tests that A. flavus, A.
TABLE 16. —Frequency of Aspergillus glaucus in stored grass seed and germination after 2 to 12 months' storage under
16 temperature-relative humidity (RH) conditions 12
'Each value is average for Festuca arundinacea, F. rubra, Lolium muttiflorum, and Poa pratensis; 100 seeds per sample cultured on 10-percent
sodium chloride-malt agar after sodium hypochlorite treatment; 200 seeds per sample used to determine germination.
2
Data from Kulik and Justice (1967). (With permission of Microform Internetl. Mktg. Corp. exclusive copyright license of Pergamon Press Journal
Back Files.)
TABLE 17.— Colony diameter of mass isolates of 6 Aspergillus species

grown for 24 days at 22°-24° C on malt agar substrate1 2
fumigatus, and A. niger raised the temperature of barley, oats, and

wheat from 17° to 43° C. These results were confirmed and extended by
Christensen and Gordon (1948). Mead et al. (1942) showed that the rise
in temperature can be even higher if the seed contains a high proportion
of broken kernels and cracked seedcoats. Similar hot spots can occur in
bulk seed, which contains pockets of moist seeds, as the natural in-
sulating properties of the surrounding dry seeds prevent rapid dissipa-
tion of the heat generated by respiration of the moist seeds and the
associated fungi.
Pockets of moist seeds can arise in low moisture content seeds
through roof leaks, insect activity, and moisture translocation when
temperature gradients within the seed mass are allowed to occur. The
fungi associated with hot spots in seeds stored on the farm were studied
by Wallace and Sinha (1962). They observed that hot spots may occur at
any location in a bin and that temperatures can increase to 53° C
winter. Field fungi such as Alternaria spp. were not common in the
heated seeds, although they were present in many of the viable seeds.
The heated seeds were infected chiefly by storage fungi, mainly Peni-
cillium and Aspergillus spp., especially A. flavus, A. fumigatus, and A.
versicolor. Absidia spp. were also frequently observed. Christensen
(1957) stated that "heating is likely to be the final and violent effect of
mold invasion of the seed, not an indication of beginning deterioration."
Sinha and Wallace (1965) investigated the succession of fungi and
actinomycetes populations associated with progressive stages in the
heating process of an artificially caused fungus hot spot in stored wheat.
The growth of Penicillium cyclopium and P. funiculosum in a moist
pocket of grain stored at —5° to 8° C for 4 months began the heating
process. A maximum temperature of 64° was reached in this hot spot.
Since these two fungi cannot live above 55°, bacterial and biochemical
activity was deemed responsible for the increase to 64°. The following
ecological succession of micro-organisms, which often overlapped, was
observed: Penicillium cyclopium, P. funiculosum, Aspergillus flavus,
A. versicolor, Absidia spp., and Streptomyces spp.
Several investigators have studied the mycoflora of freshly harvested
and stored peanuts. Hardin (1966) observed the internally borne my-
coflora of Georgia peanuts . Fusarium spp. were found in 20.7 percent,
Penicillium spp. in 18.5 percent, and Aspergillus spp. In 7.5 percent of
the seed at harvest. Jackson (1965) used dilution methods to investigate
the mycoflora of soil that adhered to dry peanut pods. Relatively small
numbers of Aspergillus flans, A. niger, A. terreus, Rhizopus spp., and
Sclerotium bataticola were found. Penicillium citrinum, P. funiculo-
sum, P. rubrum, and Fusarium spp. were found in large numbers.
Aspergillus flavus, A. niger, Rhizopus spp., and S. bataticola ex-
tensively penetrated pods and kernels when dry; the infested pods were
hydrated for 6 days at 26°, 32°, or 38° C. As the temperature increased,

infection by A. flavus and A. niger increased.
Garen (1966) studied the endogeocarpic floras of Virginia peanuts and
reported that. "Trichoderma viride seems a dominant and Penicillium
spp. seem sub-dominants in the climax endogeocarpic community of
sound and rotting peanut pods; and Aspergillus flavus and A. niger,
which have a potential for causing trouble, are quantitatively minor but
persistent species in this flora."
Diener (1960) investigated Georgia farmers' stock peanuts (uncleaned
and unshelled) that had been stored for 8 to 56 months. He found that
the predominant fungi consisted of certain species of the Aspergillus
glaucus group (amstelodami, chevalieri, repens, and ruber), A. res-
trictus, A. tamarii, Cladosporium spp., Penicillium citrinum, Torula
sacchari, and members of the Mucorales. Large numbers of fungi were
directly associated with kernel moistures of 12.5 percent or greater at
the time the seeds were stored. Kernel moisture content decreased to
about 7 percent, which is considered safe, in 3 to 4 weeks after the bins
were filled, regardless of the initial moisture content of the kernels.
During the 5-year storage period the moisture content of no sample
ever exceeded 6.8 percent. There was no relationship between per-
centage of initial kernel damage, final damage, or increased damage
during storage.
Control of Fungal Deterioration

Deterioration of stored seeds by fungi is controlled principally by
drying the seeds to a safe moisture content prior to storage in a dry
place. Most storage fungi cannot invade seeds that are in moisture
equilibrium with 65-percent relative humidity or lower. However, some
fungi are xerophytic and can invade seeds that contain only slightly too
much moisture.
Because the moisture content of a seed lot represents the average of
many seeds, some individual seeds may possibly contain an unsafe
amount of moisture even though the moisture content of the lot is at a
safe level. Seeds containing too much moisture are susceptible to
invasion by storage fungi, which may then spread throughout the lot.
Therefore it is extremely important to accurately measure seed mois-
ture content. For best results, seeds should be dried as soon as possible
after harvest, then thoroughly cleaned prior to storage in an environ-
ment where they will not absorb moisture.
Fumigation to control insects ordinarily has no detectable effect on
storage fungi (Christensen and Hodson, 1960).
For the effects of micro-organisms on stored cereal seeds, see Se-
meniuk (1954), Christensen and Kaufmann (1969, 1974), and Wallace
(1973).
Do not assume that the effects of storage fungi on seeds of other field
crops, flowers, or vegetables will necessarily be the same as those on
stored cereal seeds. For example, KuIlk (1973) found that seeds of
cabbage, cucumber, pepper, radish, and turnip inoculated with spores
of either of two common storage fungi and stored at 85-percent relative
humidity and 22°-25° C for 30 days remained largely free of invasion.
Qasem and Christensen (1958) found that 84 percent of the corn seeds
they inoculated with a storage fungus and stored at 85-percent relative
humidity and 20° were invaded by the fungus after 1 month. Also, Tuite
and Christensen (1957) found that 100 percent of the wheat seeds they
inoculated with a storage fungus and stored at 80-percent relative
humidity and 25° were infested after 1 month.
Insects
As with storage fungi, much of our knowledge of insects that attack
stored seeds comes from studies of stored grain and grain products.
According to Henderson and Christensen (1961), stored seeds are
attacked primarily by those insects that destroy stored grain and grain
products. Of the several hundred insect species associated with stored
grains and seeds, only about 50 cause problems. Of that 50 only about a
dozen cause serious damage, including the rice weevil, granary weevil,
lesser grain borer, Angoumois grain moth, cadelle, sawtoothed grain
beetle, khapra beetle, and flat grain beetle. The weevils puncture the
seedcoat and destroy the endosperm, but other stored-product insects
attack the embryo. In either case the germination potential is reduced
or totally destroyed.
Stored-product insects that fly migrate to previously noninfested
seed storage areas. Seeds may also become infested from contaminated
storage bins or sacks or in the field prior to harvest. One species of
chalcid lays its eggs individually in seeds of alfalfa, certain clovers, and
trefoils. The larva gradually consumes the contents of the seed and
becomes an adult wasp. It then chews its way out of the seed, mates,
and the female lays her eggs in host seeds, whether on the plant or in a
seed warehouse.
Control measures recommended by Lieberman et al. (1961) are
designed to destroy as many as passible of the larva-infested seeds:
"One should clean all seed carefully and destroy or use the cleanings,
prevent seed from forming on volunteer plants, clear the field after
harvest, work all chaff into the sell, and, if necessary, provide moisture
to encourage the growth of fungi present in the soil that will kill the
overwintering larvae in the seeds."
Additional insect control measures include thorough cleaning and
fumigation of all seed-handling equipment, seed containers, and storage
areas, and when required an insecticide application.
For some types of insects, fumigating the seeds before or during

storage is also essential to good control. Seed fumigation must be very
carefully done, as some fumigants under certain conditions are very
harmful to seed germination. Methyl bromide is especially harmful to
seed germination when improperly applied.
Cotton (1954, 1960) reported on the role of insects in deterioration of
stored seeds.
Rodents
Rats, mice, and squirrels destroy thousands of pounds of seed each
year. Much of the loss is not what these rodents eat, but what they
scatter and mix. They have been reported to destroy as much as 200
million bushels of grain each year in the United States (Cotton, 1960).
Even greater losses occur in other parts of the world. The best control
for rodents is to keep them out of seed storage areas. This can be
accomplished by rodentproof construction and by eliminating such
entryways as cracks and holes in walls and floors and unscreened
ventilators. Cleanliness inside and outside storage areas also helps to
keep rodents away.
Rodent infestations can be eliminated by using traps, poison baits, or
fumigation. Generally toxic gases are handled by an expert. Spencer
(1954) and Cotton (1960) published detailed accounts of rodents in grain
and their control.
Fungicides and Fumigants

Seed treatments have long been recognized as injuring; seeds in
storage. Clayton (1931) reported that the recommended treatment for
many kinds of seeds with mercuric chloride, liquid organic mercurials,
or hot water greatly reduced seed longevity. However, seeds treated
with fungicide dusts usually suffered no injury from the treatment.
Crosier (1938) found abnormal germination in wheat seeds that had
been treated with Ceresan and held in cool, dry storage for 1 year prior
to being tested for viability. Corn seed of 12-percent moisture or less
was treated by Koehler (1938) with several mercurials and stored up to
1 year. When treated seed was stored near the roof of an open shed, it
was definitely injured every time. In general, in this study, freshly
treated seeds performed better by a small margin.
Miles (1939) showed that storage of cottonseed treated with Ceresan
or New Improved Ceresan in a dry laboratory up to 17 months caused
no injury to the seed. In another study, Miles (1941) stored mechani-
cally delinted cottonseed treated with New Improved Ceresan in 100 -
pound bags (storage conditions nut given). Germination tests showed no
injurious effects after 17 months in cotton bags.
Seeds of alfalfa, ladino clover, red clover, and sudangrass were

treated with New Improved Ceresan, Arasan, Spergon, Semesan, or
yellow cu.procide by Kreitlow and Garber (1946) and were stored in
closed or open containers at 10° or 25° C for 30 months. At 25°, only the
germination of sudangrass treated with New Improved Ceresan was
greatly reduced. At 10°, germination of treated seeds and controls
stored in closed containers was much better than that of seeds stored in
open containers. Jute seeds treated with a mercuric acetate or ethyl
mercuric chloride dust were stored by Ghosh et al. (1951) for more than
14 months in airtight, moisture-free, nonporous containers without any
reduction in viability.
Koehler and Bever (1954) reported reduced germination of oats and
wheat caused by volatile mercury compounds. The reduction increased
as dosage, length of storage, and moisture content of the seed in-
creased. Compounds differed in the amount of injury they caused.
Reduced viability was not always related to volatility of the compound.
In another study, wheat seeds of two cultivars of about 10-percent
moisture content were treated by Koehler and Bever (1956) with 11
organic mercury fungicides and stored for 10 months in stoppered
bottles at 4.5° or 21° C. Germination tests in sand showed that most of
the treatments caused significantly lower stands than in the nontreated
controls. Also, there were striking differences among the compounds in
the amount of injury they caused. In general, however, they found that
seed storage at low temperature resulted in less seed injury.
Nakamura et al. (1972) treated seeds of beet, cabbage, carrot, egg-
plant, pea, pepper, and tomato with four organic mercury and four
organic sulfur fungicide dusts. The seeds were then stored for up to 3
years. Organic mercury compounds greatly reduced the viability of
eggplant seeds and caused some injury to seeds of beet and cabbage.
Seeds of carrot, pea, pepper, and tomato were not injured by these
compounds. Seeds of these vegetables treated with organic sulfur dusts
and stored for 1 year did not undergo reduced germination. Properly
applied fungicides usually do not impair germination during favorable
storage conditions.
Young (1929) investigated the effect of several fumigants on seed
viability for controlling rice weevils. He found that seeds of alfalfa,
barley, beans, buckwheat, clover, corn, cowpeas, lima beans, oats, rye,
sunflower, timothy, and wheat could be treated with up to twice the
minimum lethal concentration of ethylene dichloride, isopropyl formate,
tertiary butyl chloride, and trichloroethylene without seriously reduc-
ing their viability. Ethylene oxide and methyl chloroacetate seriously
reduced germination.
Cobb (1958) fumigated seeds of 14 field crops and 15 vegetable crops
at 3 moisture contents with methyl bromide. Germination tests were
made immediately after fumigation and after storage for up to 90 days.

Some seeds were stored for up to 2 years. He found that injury
occurred while seeds were in contact with the methyl bromide, that
fumigant injury was not reversible, that seeds uninjured by fumigation
did not exhibit subsequent injury, and that some seeds injured by
fumigation declined in viability more than the nontreated control seeds.
Based on his study, Cobb recommended that "the fumigation of dry
seed at low temperatures and gas concentrations offers the greatest
opportunity for the safe use of methyl bromide." He also recommended
"that for fumigation when seed viability is important, test fumigation
and germination should be done before large-scale fumigation is at-
tempted with methyl bromide."
King et al. (1960) evaluated hydrogen cyanide, methyl bromide, and
carbon tetrachloride alone and in mixtures with several other fumigants
for their effect on germination of barley, corn, cotton, oats, rice,
sorghum, and wheat at three seed moisture levels and three treatment
temperatures immediately after fumigation and after storage in poly-
ethylene bags for 12 months at room temperature. They found that
either a high seed moisture content or a high temperature during
fumigation caused a reduction in viability. A high seed moisture content
combined with a high temperature usually resulted in extensive injury.
Fumigated seeds of all seven crops produced less vigorous seedlings
than did the nonfumigated controls, although in some cases differences
in germination percentages between the fumigated and nonfumigated
seeds were minor. The rate of germination of stored seeds was de-
creased by fumigation. The authors recommended that "germination
tests should be performed immediately before sale if fumigated seed
have been in storage for a prolonged period."
Potgieter and de Beer (1972) treated white corn, yellow corn, and
grain sorghum with methyl bromide gas or phosphine gas, then planted
the seed in a sandy loam soil after about 13/8 inches of rainfall. Methyl
bromide fumigation was rather injurious and repeated fumigation with
phosphine decreased field performance.
CHANGES ASSOCIATED
WITH SEED DETERIORATION
Biochemical Changes
Seed storage begins immediately after maturity regardless of where
or how seeds are held. During seed development anabolic processes
predominate and bring about a gradual increase in dry matter, including
development of an embryo and food reserves. Following maturation
biochemical changes continue and eventually catabolic processes pre-
dominate and deterioration becomes apparent. Catabolic changes occur

more slowly under low temperature and low relative humidity than
under high temperature and high relative humidity.
Although much has been written about chemical changes associated
with seed deterioration (Crocker and Barton, 1953; Zeleny in Anderson
and Alcock, 1954; Abdul-Baki and Anderson in Kozlowski, 1972; Rob-
erts, 1972), our knowledge of biochemical deterioration of seeds is still
inconclusive. Attempts have been made to correlate such factors as
depleted food reserves, increased enzyme activity, increased fat acid-
ity, membrane permeability, and similar changes with deterioration.
Biochemical changes in the aging process take place in both the embryo
and endosperm. By embryo transplant technique Kikuchi (1954) and
Floris (1970) found that embryos from new seeds grew slower when
transplanted to old ("decrepit") endosperm than when transplanted to
the endosperm of new seeds. Kikuchi (1954) reported that the embryo
deteriorated more rapidly than the endosperm, and he theorized this
was because of fundamental differences in their nature. Floris (1970)
concluded that aging is a progressive process accompanied by accumu-
lation of toxic metabolites, which progressively depress germination
and growth of seedlings with increased age. Obviously important bio-
chemical changes take place in the embryo and endosperm during
storage, but the significance of these changes in relation to deterioration
is not fully understood.
Villers (in Heydecker, 1973) proposed that deterioration in dry stored
seed results from the lack of operable systems to repair and replace
organelles, whereas in moist stored seeds repair systems are fully
operable. He pointed out that the full consequences of damage accu-
mulating during storage are not evident until imbibition.
Germination and Vigor

From prehistoric times man has known that the germination capacity
of seeds declined with age (James, 1967). Even today most studies of
physiological and biochemical changes in seeds, oilseeds, and grain
include plans for determining the percent germination as a criterion of
deterioration or change.
Vigor is considered here in relation to the seed's strength or power of
germination, its ability to send out a strong root and shoot even under
conditions of stress, and its freedom from attack by micro-organisms.
Loss in vigor can be thought of as an intermediate stage in the life of
the seed, occurring between the onset and termination of decline
(death). Decline in vigor is extremely difficult to measure. No generally
accepted method has yet been found to measure vigor, but the most
useful available methods are based on measurements relating to ger-
mination capacity. For example, Abdalla and Roberts (1969b), working

on seed deterioration of barley, broadbeans, and peas, reported that the
percentage of seed viability is an excellent indicator of growth potential
of the surviving seeds, irrespective of the factors responsible for the
loss of viability. Gill and Delouche (1973) found rate of germination and
seedling growth of corn to be the most consistent and sensitive meas-
ures of the progress of deterioration. A similar conclusion was drawn
by Christidis (1954), who found that the rate of germination of stored
cottonseed increased for the first 5 years of storage and then gradually
decreased throughout the following 6 years.
As the catabolic changes continue with increasing age, the ability of
the seed to germinate is reduced. The curve in figure 2 shows that
measurable decline in viability or germination capacity does not begin
immediately after maturity. Under favorable storage conditions the
initiation of decline may be a few months to many years depending on
storage conditions, kind of seed, and previous storage conditions. When
making germination tests on stored seed, one of the first indications of
deterioration is reduced vigor, which is shown by reduced rate of
germination and production of weak or watery seedlings and seedlings
with stunted radicles. Seeds with reduced vigor would appear to pro-
duce low yields compared with fresh vigorous seeds; however, this is
not always true.
Yield
Old seed can be vigorous and alternatively new seed can be de-
teriorated depending on the storage conditions to which they have been
subjected. Therefore seed of a given chronological age may or may not
be deteriorated. Some of the pertinent literature does not make this
distinction. If a maximum yield of a given crop is to be obtained, a
minimum number of plants per acre (or hectare) is necessary. If the rate
of planting low germinating seed is increased to satisfy this minimum
requirement, yield is not decreased compared with the yield from new,
vigorous seed. The problem is whether plants from seeds of low vigor
can compete in the field with plants from vigorous seeds.
Abdalla and Roberts (1969a) planted seeds of barley, broadbeans, and
peas in a culture medium and in the field. Plants from some seeds
associated with decreased viability exhibited retarded growth of roots
and shoots and increased variability in growth during early life. The
early inhibition of growth rate did not persist, and there was evidence
that under normal cultural conditions the low growth rate during early
plant development might be compensated for in later growth stages.
They concluded that unless the germination of deteriorated seed of
barley, broadbeans, and peas is less than 50 percent, the final yields are
not significantly affected.
Christidis (1954) found that plants from cottonseed up to 10 years old

yielded about the same weight of seed cotton as plants from fresh seed.
Others who have reached similar conclusions from work with seeds of
different crop species are Rodrigo (1939) with mung beans, Barton and
Garman (1946) with aster, lettuce, pepper, and tomato, Fulutowicz and
Bejnar (1954) with sugar beet, and Ghosh and Basak (1958) with jute.
Barton and Garman (1946) found yields of crops from seeds of the four
species, which had been stored at —5° C up to 13 years, were equal to
those of fresh seeds. Thirteen-year-old lettuce seed produced larger
heads than did fresh seeds. Likewise plants from the 11-year-old mung
bean seed used by Rodrigo (1939) produced significantly higher yields of
pods and beans than plants grown from fresh seed.
Not all the evidence is in favor of normal yields of crops produced
from deteriorating seed. Newhall and Hoff (1960) stored onion seeds for
22 years over calcium chloride in a closed desiccator. Field tests
revealed a 30-percent loss in yield by the 22-year-old seed compared
with yield of fresh seed. Bulbing vigor was reduced by 15 percent,
although this was hardly detectable by casual observation.
Chirkovskiy (1953) reported that longtime storage of tobacco seed
resulted in loss of germination and lower yields of leaves per plant
compared with yields of plants from fresh seeds. He (1956) indicated
that aging of seeds had a depressing effect on plant development,
especially during early growth and that unfavorable storage conditions
increased the depressing effect.
Corn that germinated only 38 percent in the field yielded only 50
percent as much grain as seed that germinated 80 percent in the field
(Heft and Munn, 194?). It is not clear whether the reduced yield was due
to reduced field stand, weakened vigor, or both. They also claimed
reduced yields from aging seeds of cucumber and soybean. Presumably
they credited the reduced yields to low seed vigor, as seeds of both
crops produced seedlings with stubby roots insufficient to properly
anchor the plant or to provide an adequate supply of soil nutrients.
Because information is limited on plant yield from seeds of low or
weakened vigor, future research should segregate the principal factors
so the contribution of each can be ascertained. Among these factors are
use of uniform genetic stocks, isolation of low vigor effects, and effects
of competition versus noncompetition with other plants under normal
field conditions.
Cytological Changes
One of the changes associated with seed aging is aberration of
chromosomes, sometimes referred to as mutagenic effects. tie Vries is
credited with discovering mutations in aging seeds in 1901 (Kostoff,
1935). Some of the chromosome alterations in aged seed include frag-

mentation, bridges, fusion, ring formation of chromosomes, and varia-
tion in nuclei size. These changes have been found in a relatively large
number of crop and native plant species, including Antirrhinum,
Crepis, Datura, Nicotiana, barley, broadbean, corn, onion, pea, rye,
sugar beet, and wheat.
According to James (1967), there is no evidence that these cytological
aberrations develop in seeds stored under favorable conditions. The
seeds used in most experimental work so far have been subjected to
high storage temperatures or humidities or both. Possibly James'
statement can be questioned depending on what is considered favorable
or unfavorable storage conditions. For example, Blakeslee (1954) found
mutations in Datura seeds stored at room temperature. He did not
indicate that either the temperature or the relative humidity was
unfavorable during storage. He also found a very low frequency of
mutations in Datura seeds that had been buried in the soil for 39 years.
HOW SEEDS ARE DRIED

Characteristics of Water
Because drying involves removing water from the seed, let us first
consider some of the characteristics of water. In nature, water exists
simultaneously as a solid, liquid, and gas. Water freezes at 0° and boils
at 100° C. It contracts as it cools to near the freezing point but expands
on freezing. The quantity of solid or liquid water is expressed by weight
or volume, whereas the quantity of gaseous water is given as percent
humidity, either relative or absolute. The amount of water in a seed is
its moisture content, which is measured on a wet or dry weight basis
with respect to the seed weight. When seed moisture content is given on
the wet weight basis, the amount of water in the seed is a percentage of
the seed weight before the water is removed. When seed moisture
content is expressed on the dry weight basis, it is a percentage of the
seed weight after the water is removed.
During development, maturation, and ripening, the water content of
seeds gradually decreases until harvested seeds finally dry to the point
where there is no further reduction in moisture because their water
content reaches equilibrium with the ambient relative humidity. Any
further change in seed moisture content will be the result of a change in
either the relative humidity or the temperature of the air or both.
Because of chemical differences in their composition, not all kinds of
seeds will equilibrate at the same moisture content at a given relative
humidity. In fact, not all cultivars of a given kind of seed nor even all
lots of a cultivar have exactly the same equilibrium moisture content for
any given relative humidity.
Water in seeds is present in liquid form in cell walls and cell contents
and in gaseous form in intercellular spaces. Since water is removed
from seeds by evaporation, it is removed only in its gaseous form.
Principles of Drying
Seed drying requires the transfer of heat, because a seed can be dried
only by evaporating moisture from its surface, and the heat content of
moisture vapor is greater than that of liquid water. Heat transfer may
be accomplished by contact, convection, or radiation. Most seed-drying
equipment transfers heat by convection. Drying of a seed requires that
evaporation of moisture from its surface be accompanied by the
transfer of moisture from its interior to its surface. When water
evaporates from the seed surface into the atmosphere, a moisture
gradient is set up inside the seed that causes internal moisture to move
toward the surface. If evaporation from the seed's surface occurs too
rapidly, the extreme moisture stress that develops can, and usually
does, damage the embryo and cause loss of viability. It is therefore
essential that seed drying be carefully controlled to prevent stress
damage. It is also important that the vapor pressure of the surrounding
atmosphere not be allowed to increase. If the vapor pressure of the
atmosphere becomes greater than that of the seed surface, the seed will
gain rather than lose moisture. Additional information on principles of
drying seeds can be found in Anderson and Alcock (1954), Hall (1957),
Brandenburg et al. (1961), Kreyger (1963a), Harrington and Douglas
(1970), and Philpot (1976).
Methods of Drying Seeds

Methods of drying seeds include natural drying, sun drying, un-
heated, heated, and dehumidified air-drying, drying in storage and
before storage, drying with desiccants, vacuum drying, and freeze
drying. These methods may be used alone or in various combinations.
Natural Drying
Natural drying occurs in the field during ripening and in the bin after
harvest. It is the process by which seeds lose water naturally without
help from man. The extent of natural drying is regulated by such
factors as air temperature, relative humidity, and wind velocity. Shat-
ter resistance, harvesting procedures, and workload of the producer
also affect the amount of natural drying permitted before harvest. A
hot, dry wind prior to full maturity can seriously damage a seed crop by
causing too rapid drying, but a moderate temperature and relative
humidity can produce top-quality seed. Some but not all seed crops are
harvested or partially harvested before the seeds dry to moisture
equilibrium with the atmosphere. Such seeds must receive additional

drying. Sometimes the plants are cut and placed in windrows and this
speeds up natural drying.
Sun Drying
Where sun drying is possible, seeds may be spread in screen-bottom,
trays placed on low racks in the sun or on flat-roofed houses, or they
may be spread on the ground or on concrete. Seeds dried in the sun are
stirred from time to time to facilitate rapid, uniform drying. Sun drying
is used for such seeds as cantaloup, cucumber, pumpkin, tomato,
watermelon, and others that are threshed wet. Because these seeds do
not separate readily from the surrounding pulp, fermentation, chemical
treatment, or mechanical action is used to help free them. Once freed,
they are washed thoroughly and placed in screen-bottom trays to be
cured in the sun. Although these and other kinds of seed are frequently
dried naturally, they may also be dried artificially. Most seeds har-
vested at a moisture content too high for safe storage are dried
artificially. Solar heat can be used in the bin drying process (fig. 7).
Artificial Drying
Three basic types of seed dryers are layer, batch, and continuous
flow. There are sound engineering and economic reasons why each type
is better suited for certain kinds of drying than the others. No one type
of dryer is best suited for all drying needs. The user must choose the
best one for his specific needs. One should not attempt to select a seed
dryer without first obtaining sufficient information on the advantages,
disadvantages, and limitations of all available dryer systems. One must
also have sufficient information on the specific drying requirements of
the kind or kinds of seeds to be dried. Table 18 presents data for several
crops. Although not complete, the data provide a reference for related
kinds of seeds.
Layer Drying.—Layer drying is usually in-storage drying, which
may be done with or without supplemental heat. For layer drying, the
storage area is equipped with a network of air ducts attached to a large
fan. The air ducts (fig. 8) should be placed on the floor of flat storage
structures so they provide reasonably uniform airflow, cooling, and
drying in the entire mass of seeds. The manifold should be as close to
the wall as possible, and in round buildings it should be opposite the
door so as not to interfere with unloading. The method of unloading
must always be considered when locating the aeration tubing. Other
factors such as depth of seeds, accumulation of fine particles, and
uneven loading have a direct bearing on operation efficiency of the
drying system. The size and speed of the fan are determined by the size
of the storage area, the kind and depth of seed to be dried, the
TABLE 18. —Static pressures and estimated quantities of various seeds

and grains that can be dried per batch per fan horsepower for
different airflows and seed depths1
TABLE 18.--Static pressures and estimated quantities of various seeds

different airflows and seed depths1—Continued
TABLE 18.—Static pressures and estimated quantities of various seeds

different airflows and seed depthsl—Continued
temperature of the air, and the amount of air movement desired. Each
layer of seed is partially dried before the next is added. The rate at
which the dryer can be filled depends on the amount of moisture to be
removed and how rapidly it is removed. The entire depth of seed is
finally dried in place. Because the dried seeds remain in the dryer, this
type of dryer is not satisfactory for large operations or for those that
require a variety of drying conditions.
FIGURE 8.—Diagram of air duct arrangement for square, round, and rectangular
in-storage seed and grain dryers.
Harvesting for storage in layer dryers can be accelerated by using

several bins, supplemental heat, or both (fig. 9). However, the amount
of heat used must not overdry the lower layers of seed before the top
layers are dried. One cannot simply turn up the heat to dry faster
because this may seriously overdry the lower layers. According to
McKenzie (1966), layer drying, contrary to popular belief, requires
superior management. Layer drying is suitable for on-the-farm drying
of corn, soybean, and cereal grain seeds.
Batch Dryers.—Batch dryers may be either portable or stationary.
Portable batch dryers may or may not have a burner attached. Some
batch dryers are specially designed wagons with porous beds that
permit heated or unheated air be forced upward through the seeds,
whereas others are simply drying chambers. The wagon-type dryer can
be taken to the field for filling, whereas the other types cannot. in any
case, the batch-type dryer can dry only a fixed amount of seed at a time.
After drying, the seed has to be cooled before it can be transferred to
storage. Because of the time involved in drying and cooling, it is
necessary to have several drying chambers for efficient harvesting and
drying of seed crops. Since stationary batch dryers usually have a
FIGURE 9.—In-bin layer drying with supplemental heat. Bins are filled in rotation, one
layer at a time. As each layer partially dries, successive layers are added until
storage is full and the grain is dried in place. (Courtesy of Bruce McKenzie, Purdue
Univ.)
greater capacity than portable dryers, they are preferable for com-
mercial installations.
Stationary batch dryers may be a bin, column, or rotary drum. In
bin dryers the seeds are spread over a perforated floor, and in a column
dryer they stand in a vertical column. Depth of seeds over the floor or in
the column varies with the type of seed being dried. In the rotary
dryers a revolving, perforated drum slowly tumbles the seeds as heated
air passes through them.
Batch-type dryer installations vary from a single drying chamber to
many such chambers. Batch-type drying plants generally utilize one of
five drying systems: (1) Double or two pass, (2) single-pass reversing,
(3) single pass, (4) suction, or (5) tunnel. Batch dryers are suitable for
both on and off the farm drying of many kinds of seeds, including cereal
grains, corn, millet, sorghum, and soybean.
In the double- or two-pass drying system (fig. 10), the heated drying
air is first directed through a bin containing nearly dry seeds, where the
air picks up a small amount of moisture and loses a little of its heat. The
air is than transferred and exhausted through a bin of high moisture
content seeds, which need warming up. The double pass makes a more
FIGURE 10. —Double- or two-pass drying system. (Courtesy of Corn States Hybrid Serv.,
Des Moines, Iowa.)
complicated drying system; however, it adds about 25 percent to the

drying capacity of a drying plant and reduces fuel costs substantially,
although power costs are increased some. The two-pass system pro-
vides added protection to the viability of the seeds by using cooler air
(27°— 32° C) on the higher moisture seeds. Only the relatively dry seeds
are subjected to a possible 43°.
For the single-pass reversing drying system (fig. 11), the seed drying
plant is designed to allow a single pass of air with provisions for
reversing the direction of airflow through the various bins to facilitate
uniform drying. In the single-pass system the air goes through the seed
and is exhausted. During the early stages of drying, a great deal of heat
is transferred from the air to the seeds and considerable water is taken
up by the air; however, toward the end of the drying cycle, little heat or
moisture is transferred, resulting in considerable drying potential being
wasted. This method has approximately 80 percent of the drying
efficiency of the two-pass system.
The single-pass system (fig. 11) has no provision for reversing air-
flow; consequently, it has only about 75 percent of the efficiency of the
two-pass system. With this system the seeds on the bottom of the bin
will be overdried and those on the top underdried. A uniform moisture
content is obtained by blending after the seeds are removed from the
dryer.
In the suction drying system (fig. 12) the blower is placed at the
exhaust side or the drying bins. This system can be used with either the
FIGURE 11.—Single-pass and single-pass reversing drying system. (Courtesy of Corn

States Hybrid Serv., Des Moines, Iowa.)
FIGURE 12.—Suction two-pass drying system. (Courtesy of Corn States Hybrid Serv.,
Des Moines, Iowa.)
one-pass or two-pass design and is worthy of careful consideration when

a new installation is planned.
The tunnel dryer is another form of the heated air batch dryer. It is
used for certain vegetable seeds, such as cucumber, muskmelon, and
tomato, and for drying seeds in bags. Because vine crop and tomato
seeds have to be washed to remove adhering fruit pulp, they require

rapid drying immediately after washing. Rapid drying is accomplished
by spreading the seeds in thin layers in screen-bottom trays, which are
either placed in a tunnel dryer or dried in the sun, as explained
previously. Tunnel dryers consist of open trenches through which
heated air is circulated up through the seeds in the trays. Seeds in bags
can be dried in a tunnel dryer by building the tunnel with a cover having
special holes over which the bags of seeds are placed. This is the most
inefficient system of drying seeds. A tunnel dryer can be constructed
with a capacity to meet a specific need.
Continuous Flow Dryers.—The continuous flow dryer may be hori-
zontal or vertical (fig. 13). It is called continuous flow because the seeds
move slowly from the point of entry to the point of exit and are dried as
they move through the unit. A continuous flow dryer is not suited for
drying small quantities. For best efficiency, the system must be full of
seeds at all times. Consequently, the continuous flow dryer is not at all
suited to an operation requiring frequent changes from one kind or
cultivar of seed to another. Its greatest value is for grain drying where
thousands of bushels of a single cultivar can be dried without interrup-
tion.
Modification of Basic Dryer System.—The dryeration (fig. 14) repre-
sents a new grain drying process developed at Purdue University. It is
a combination of high-speed and high-temperature drying and slow
cooling (McKenzie et al., 1968). With the dryeration process, no cooling
is done in the dryer. Grain is discharged at 50° to 60° C from the dryer,
still carrying 1- to 3-percent excess moisture into a separate cooling bin.
The hot grain is held without mechanical cooling for a tempering period.
It is then cooled slowly to remove the excess moisture before being
transferred, dry and cool, to storage or load out. The dryeration
process, because of the high drying temperature and the tempering
period, may not be as satisfactory for seed drying as it is for grain
drying.
Dehumidified Air Systems.—In dehumidified air systems, a dehumi-
difier dries the air before it is heated and passed through the seeds. The
air dryer converts the latent heat in the moisture to sensible heat in the
drying airstream, which lowers the relative humidity of the air by
decreasing the total moisture in the air. The rise in temperature
associated with dehydration reduces the amount of heat needed from
the main heat source, which reduces fuel costs while drying efficiency is
increased. Dehumidified air can be used with either a batch-type or a
continuous flow dryer. All that is required is to install a dehumidifier in
the air delivery system.
Dehumidification, although generally used with a closed system , can
be incorporated into any drying system. The air passes through the
FIGURE 13.—Vertical continuous flow dryer. (Courtesy of Ferrell-Ross, Oklahoma City,

Okla.)
dehumidification medium, usually silica gel, and then is circulated

through the rest of the system back to the dehumidifier (fig. 15). Since
the drying medium has to be recharged or dried out, either a two-tower
or a rotary system is used. With the two-tower system (fig. 16), one
tower dehydrates the air while the drying medium in the other tower is
being dried. The switch from one tower to the other is automatic. With
a rotary dehumidifier (figs. 17, 18) the drying agent is held in a circular
unit, which continuously rotates so that one section is drying the air
while the desiccant in another section is being dried.
FIGURE 14.—Schematic flow diagram of dryeration process. (Courtesy of Coop. Ext.

Serv., Purdue Univ.)
In general, seed drying plants are custom engineered for specific

purposes (figs. 19, 20). For example, a dryer plant designed to dry ear
corn cannot be used to dry Kentucky bluegrass strippings. Dryers, as
well as drying plants, are sometimes designed for special purposes. A
special-purpose dryer (fig. 21) was developed by the National Seed
Storage Laboratory for conditioning small quantities of seed for storage
research projects. The trays used in the dryer were designed for loose
seed; however, by means of a system of cross wires to allow air
movement between packets, the trays can be used for either bulk or
packet drying, especially if the wire grid is removable.
A special walk-in seed-drying room was constructed at the New York
Agricultural Experiment Station to handle relatively large samples of
plant materials in bags (fig. 22) (Dolan et al., 1973).
Miscellaneous Dryers.—Seeds can also be dried under a partial
vacuum, by the freeze-drying process, or by infrared radiation
(Schroeder and Rosberg, 1959), all of which are used extensively in the
food industry. Each of these drying methods has certain advantages
and disadvantages. In general, they are more costly than conventional
drying methods, and in some cases they are suitable only for drying
small quantities of seed. In vacuum and freeze-drying, seed moisture
content can be reduced to very low levels without using excessively high
temperatures. Seed dried to moisture levels below air dryness must be
stored in either moisture-barrier containers or dehumidified rooms;
otherwise the advantage of such drying will soon be lost through
rehydration.
Much of this information was reported by Anderson and Alcock
(1954), Hall (1957), Wheeler and Hill (1957), Brandenburg et al. (1961),
Kreyger (1963a, 1963b, 1972), Barre (1963), McKenzie (1966), McKenzie
et al. (1968), Harrington and Douglas (1970), Peterson (1971), and

Philpot (1976).
Factors To Consider in Selecting a Dryer System
Drying Capacity.—The drying capacity should be sufficiently large to
permit continuous, efficient harvest of the seed crop, regardless of
FIGURE 15. —Flow diagrams of air movement through a combination dehumidified-air

heated-air dryer system. (Courtesy of Corn States Hybrid Serv., Des Moines, Iowa.)
whether only one or several cultivars are to be dried. Daily capacity of

the drying system should efficiently handle one day's harvest.
investment Necessary To Get Capacity.—If the investment in drying
capacity is excessively high, costs of drying are bound to be high. Since
ownership costs are often about half the cost of drying, the initial cost
as well as the maintenance costs should be carefully considered when
selecting a dryer.
Fuel and Power Costs.—The cost of fuel to heat the drying air and
FIGURE 16.—Two-tower dehumidifier for use in storehouses. (Courtesy of Seedburo

Equipment Co., Chicago, Ill.)
power to operate the fan may not vary much among dryers; however,
such costs must be included when total cost of drying is calculated.
Since high-speed systems tend to increase costs more than capacity,
they cost more to operate.
Airflow.—Efficiency of a dryer is directly related to airflow, which is
determined by the output of the fan. Generally the airflow per horse-
power does not vary greatly among fans of comparable size. However,
some industry rating practices are confusing. For example, fan motors
sometimes deliver more horsepower than their nameplate rating.
Nameplates on fan motors sometimes carry a dual rating, showing a
range of horsepower. Such practices tend to make one fan seem more
efficient than another with the same horsepower rating when the
difference is actually the result of one motor delivering a higher
horsepower.
FIGURE 17. —Cutaway view of a rotary dehumidifier. (Courtesy of Corn States Hybrid
Serv., Des Moines, Iowa.)
FIGURE 18 .—High capacity rotary dehumidifier with 50,000 cubic-foot-per-minute air-

flow. Equipment includes a 150,000 cubic-foot-per-minute capacity main blower and
air-heater unit. (Courtesy of Corr. States Hybrid Serv., Des Moines, Iowa.)
FIGURE 19. --This corn drying plant utilizes the most recent advancements in drying and
handling equipment. (Courtesy of Equipment specialists, Inc., Taylorville, III.)
FIGURE 20.—The dried ear corn from figure 19 passes from the bins along the permanent
discharge conveyor directly into the sheller. (Courtesy of Equipment Specialists,
Inc., Taylorville, Ill.)
Heat Input and Drying Temperature.—Assuming the airflow is suf-

ficient for the drying method used, failure of a system is usually the
result of heat management. Systems fail because of too much or too
little heat. Too much heat reduces seed quality, and too little heat slows
drying. There are desirable and workable air-heat ratios for each dryer
system. These ratios vary with the dryer and the kind of seed. For best
quality, seed temperature should not exceed 43° C for grain and 32° for
vegetable seeds. Higher temperatures will increase drying rate but will
also reduce seed quality. Air temperature can exceed seed temperature
if the seed is not exposed long enough to reach air temperature or pass
critical limits. Table 19 gives effects of heating time, temperature, and
seed moisture content on loss of viability for several kinds of seed.
Laboratory studies by Bass (1953) and Grabe (1957) showed that the
higher the moisture content of Kentucky bluegrass and smooth brome-
grass seeds, the lower the temperature and the shorter the time at that
temperature required to bring about viability or germination loss (fig.
23) (table 20). Grabe (1957) also showed the effects of various tempera-
tures and drying periods on the germination of smooth bromegrass
seeds (table 21). He noted that none of the temperature-time treat-
ments used adversely affected the germination of smooth bromegrass
seeds, even after 29 months of storage.
FIGURE 21. —Seed dryer developed by National Seed Storage Laboratory, Fort Collins,
Colo. Above, assembled; helms', disassembled, showing twin-blower air diffuser (A),
top ventilation holes (B), and screen-bottom seed tray (C).
FIGURE 22 .—Interior of a seed-drying room, showing slotted floor, plywood covers, exit
ducts, and inlet hot air duct near ceiling. (Courtesy of N.Y. Agr. Expt. Sta.,
Geneva.)
Williams (1938) reported that it is advisable to remove excess mois-

ture from perennial ryegrass, timothy, and orchardgrass (cocksfoot)
seeds as quickly as possible after harvest. He referred to excess
moisture as being that above the norms he set. They were about 14
percent for ryegrass, 13.6 percent for timothy, and 13 percent for
orchardgrass. Because of their greater rate of absorption and slower
rate of drying, seeds of perennial ryegrass require more careful atten-
TABLE 19.—Effect of different heating times on viability of various

seeds at different seed temperatures and moisture contents (static
conditions) 1
tion during drying than those of orchardgrass and timothy. A current of

warm air at not more than 43° C alternating with a cool (15° —18°) one (20
min warm, 10 min cool) is particularly beneficial for drying ryegrass and
orchardgrass seeds. Drying with a continuous warm air current ad-
versely affected viability, even when the temperature did not exceed
35°.
Wileman and Ullstrup (1945) found that germination of ear corn with
an initial moisture content of 35 percent or more dropped rapidly when
dried at 49° C; however, corn with 25 percent or less initial moisture
suffered no appreciable germination loss when dried at 49°, whereas
corn with 20 percent or less initial moisture showed no germination loss

when dried at 54°.
Drying at too high a temperature or too rapidly can drastically reduce
seed viability (Toole and Toole, 1946; Griffith and Harrison, 1954; Raun
FIGURE 23.—Germination of Kentucky bluegrass seeds with four moisture contents after
4-48 hours at various temperatures without drying.
TABLE 20.— Viability of bromegrass seed as affected by interactions of temperature, moisture content, and duration of
treatment in 1953 and 1954 laboratory studies
TABLE 21 . —Summary of results of artificial drying experiments on germination of 13 lots of
smooth bromegrass seed in 1953 and 1954 1
1 Data from Grabe ( 1957).

2 Maximum temperatures in bottom 6 inches; temperatures in remainder of load were lower than these data.
3 Seed dried 2 hours at lower temperature, then completed at higher temperatures.
and Frey, 1957). Excessive drying can also result in loss of viability and
the development of seedling abnormalities, especially under certain
storage conditions (Evans, 1957b; Nutile, 1964a, 1964b). Heated air-
drying may induce a dormant condition, which can seriously affect field
stand establishment unless it is overcome before planting (Wright and
Kinch, 1962; Nutile and Woodstock, 1967).
Handling and Labor Requirements.—The various drying methods
obviously differ in the amount of handling equipment and labor needed.
In-storage layer drying involves two seed handling operations—filling
and unloading the bin. Batch and continuous flow drying require four
operations, namely, loading and unloading the dryer and filling and
unloading the bin. The volume of seed to be dried determines handling
equipment cost rather than the type of dryer. A large volume of seed
requires a more elaborate and expensive seed handling system.
Weather.—Weather is a major factor in seed drying operations. It
determines the amount of time available for harvesting and drying a
seed crop. Extended periods of cool, damp weather can markedly
increase harvesting and drying costs, whereas hot, dry winds can
reduce such costs. During periods of wet weather, extra heat and air
movement are required for efficient drying. During periods of hot, dry
weather, the amount of supplemental heat needed for rapid drying is
much less than that required during warm, humid weather.
Drying Rate.—The rate at which moisture leaves a seed depends on
how much the seed lacks moisture equilibrium with the air surrounding
it, the air temperature, and the composition, size, and shape of the seed.
When the initial seed moisture content is high, the rate is faster if the
temperature is high or if relative humidity is low. A change from very
slow to rapid air movement will increase the rate of drying. The rate of
drying drops off as the moisture content of the seed decreases. This
means that as the moisture content of seed decreases, longer exposure
to heat is required for each percentage of moisture removed.
In bulk drying the rate of moisture loss is not uniform throughout the
depth of seeds in a dryer. As the seeds dry, the interface between dry
and wet seeds gradually moves through the seed layer. Since the seeds
nearest the heated air source dry fastest and those farthest away dry
slowest, there is always a range in seed moisture levels through the
layer of seeds in a dryer.
The drying rate for individual kinds of seeds depends on their
individual properties as to release of moisture. However, various kinds
can be grouped with regard to their tendency to release moisture
(Kreyger, 1963a). There are fast drying seeds such as grasses, rape,
and sugar beet; normal drying seeds such as barley, oats, rice, rye, and
wheat; and slow drying seeds such as bean, corn, lupine, and pea (table
22).
TABLE 22.—Comparison of approximate rates of drying and releasing

moisture at 5 moisture contents for various seeds1
Williams (1938) found that by using a 20-minute warm air (43° C) and
10-minute cool air (15°-18°) alternation, moisture content of ryegrass,
timothy, and orchardgrass seeds can be reduced about 4.5 percent in
11/2-2 hours. Drying time for 25 percent or lower moisture ear corn can
be reduced about 20 percent by using 49° rather than 43° (Wileman and
Ullstrup, 1945).
The average time required to dry six lots of 21- to 25-percent
moisture ear corn to 12- to 13-percent moisture was reduced from 81 1/2
to 64 1/2 hours by raising the drying air from 43° to 49° C and thereby
decreasing its relative humidity. Drying rates for specific kinds of seed
are largely unavailable because such factors as the initial moisture
content, the temperature, relative humidity, and velocity of the air, and
the depth of the seed layer must all be considered simultaneously.
A preliminary estimate of drying rate can be obtained by recording

the moisture loss from a known weight of seed held under known drying
conditions for a specified length of time. Knowledge of whether the
kinds of seeds are slow, normal, or fast drying is very helpful in
determining the type of dryer system to install. For example, a contin-
uous flow dryer is generally suitable for quick and normal drying seeds
but not for slow drying seeds, nor is it suitable for nonfree-fiowing
seeds. Batch dryers are better for slow drying and nonfree-flowing
seeds. A low temperature is used at the start and gradually increased as
drying progresses. This practice is fairly common in the field corn
seed-producing areas where double-pass dryers are used.
Drying Seeds With Desiccants

Barrow ( 1915) reported that 5-percent calcium chloride by weight
minimized heating and deterioration of cottonseeds during storage.
Kondo and Isshiki (1936) reported that either calcium oxide or calcium
chloride could be used to desiccate rice seeds. They found that calcium
oxide absorbs only about one-third as much moisture as calcium chloride
but absorbs it faster. One kg of calcium chloride or 3 kg of calcium oxide
will absorb 1-percent moisture from 5.1 bushels of rice. According to
Kondo and Terasaka (1936), both calcium chloride and calcium oxide
desiccate rice for safe, sealed storage for food but not for seeds. Dexter
and Creighton (1948) found that wood blocks impregnated with calcium
chloride, magnesium chloride, magnesium sulfate, or sodium chloride
effectively removed moisture from stored grain; however, they did not
determine what effect, if any, storage with the impregnated wooden
blocks had on germination.
Although seeds can be dried safely with desiccants, drying large
quantities of high moisture seeds with desiccants is neither practical nor
economically feasible because of the large amount of desiccant needed
to remove 1 percent of water from each pound of seed. Use of desic-
cants with small quantities of seed in sealed containers is discussed
under "Packaging and Packaging Materials."
SEED STORAGE STRUCTURES

A considerable amount of seed becomes useless each year from
improper storage. Seeds must be stored dry and kept dry. As pre-
viously indicated, longevity of seeds is controlled primarily by seed
moisture content and storage temperature. For maximum storage life,
these two conditions must be carefully controlled. However, much of
the seed produced each year needs to be stored only from harvest until
the next planting season. Such seed may or may not require storage
under conditions other than normal air temperature and relative hu-
midity depending on the kind of seed and the local climate. A seed
AGRICULTURE HANDBOOK 506, U.S. DEPT, OF AGRICULTURE
storage facility must have certain basic features, regardless of any

adaptations for special purposes (Barre, 1954; Wheeler and Hill, 1957).
Basic Features of Storage Structures

Protection From Water
Rain, snow, ground moisture, or any source of water should never be
allowed to come in contact with seeds, as it increases seed moisture
content. High seed moisture content increases respiration, heating, and
mold growth and sometimes promotes sprouting in storage, all of which
decrease seed quality. The roof and sidewalls of seed storage structures
must be free of holes and cracks that permit the entry of rain or snow.
Cracks and knotholes in wood walls and roofs should be filled. All bolts
and screws in metal buildings should have rubber washers, especially
those in the roof. Since soil moisture can be absorbed readily on contact
by seeds, all seed storage structures must have a waterproof floor. A
wood floor should be elevated and a concrete floor should have a
moisture barrier under it. A metal floor is naturally impermeable to
water but will soon rust out if kept moist continually.
Protection From Contamination
Because individual seeds of different cultivars of various crops are
indistinguishable from each other and because the kinds of seeds are
often difficult to separate, each seed lot must be kept free of seeds from
other lots. Storage facilities should be constructed to provide maximum
protection from chance contamination. For bulk storage, a separate bin
must be provided for each cultivar. For bag storage, seeds of each
cultivar must be stacked separately. Seeds may also be stored in
toteboxes. Toteboxes and stacks of bags on pallets can be moved readily
by a forklift. All bags, toteboxes, and bins must be carefully and
conspicuously labeled.
Protection From Rodents
Since large quantities of seeds may be lost because of inadequate
protection from rodents, precautions must be taken to prevent their
entry into storage buildings. Metal and concrete buildings normally
provide good protection from rodents. With care, wood buildings can
also be constructed to keep them out. Metal bins with tight covers
provide rodent protection, and cloth bags can be treated to repel
rodents. For further information, see the section on "Effects of Pests
and Chemicals on Seed Deterioration in Storage.'
Protection From Insects
Although insects do not constitute a major problem with some kinds
of seeds, a good storage facility should be so constructed that any part,
or all of it, can be fumigated to control insects at any time. Insect

problems can be kept to a minimum by thoroughly cleaning and fumi-
gating each time a bin is emptied. Areas where bags and toteboxes are
stored should be kept free of loose seeds and trash at all times.
Cleanliness eliminates insect breeding places and facilitates insect con-
trol. For further information, see "Effects of Pests and Chemicals on
Seed Deterioration in Storage."
Protection From Fungi
Under certain circumstances, storage fungi can cause considerable
damage to stored seeds. Since fungi grow best under warm, humid
conditions, storage structures should provide cool, dry conditions.
Damage from storage fungi can be kept to a minimum by carefully
drying seeds to a safe moisture content, usually less than 12 percent,
and holding them under dry conditions. Sometimes ventilation of
storage structures, especially steel buildings, is necessary to prevent
the accumulation of translocated moisture. Temperature differentials
can cause water vapor to move from warmer to cooler areas of a bin,
usually to the upper surface. Such moisture movement can provide
conditions favorable for fungus growth unless adequate precautions are
taken to prevent its accumulation. Seeds can be treated with various
chemicals to control fungi. Fungicides are applied routinely to many
kinds of seeds as part of their normal processing. However, fungicides
to control soil fungi may not control storage fungi. For further infor-
mation, refer to "Effects of Pests and Chemicals on Seed Deterioration
in Storage."
Protection From Fire
Danger from fire loss is greatest with wood buildings; however, wood
can be chemically treated to retard burning. The hazard of a fire in wood
buildings can be reduced by cleanliness, both inside and around the
building. All seed storage buildings, especially processing areas, should
be equipped with special dustproof and sparkproof electrical outlets and
switches, which greatly reduce the chances of an electrical fire. All
wiring should be rodentproof. Although metal and concrete buildings
are fireproof, the same electrical wiring precautions should be practiced
in them as in wood buildings because dust explosions and fires can be
caused by electric sparks. Structures with these general characteristics
should provide safe storage for most kinds of seeds from harvest to the
next planting season, except in areas with extremely high temperatures
and relative humidity. In such areas, additional protective measures,
namely temperature and humidity control, are required to maintain
seed quality during storage.
Types of Storage Structures

Farm Storage
Farm storage is usually for a few days or weeks during the harvest
season. Seeds are seldom held on the farm longer than from harvest to
the next planting season. They may be stored in bins, bags, or in some
cases on the ground. Ground storage is generally temporary and used
only in an emergency. Bulk lots may be stored in wood or metal bins,
granaries, and barns. Seeds, unless very dry, keep better in wood than
in metal buildings because heat buildup is greater and moisture loss is
slower in metal than in wood buildings. Heat accumulation in metal
buildings can seriously reduce the viability of seed stored with high
moisture. Heat buildup can be reduced by painting the outside of the
building white, by ventilation, or both. Seeds in bags may be stored in
any farm building where space is available. This practice frequently
causes large losses through insect and rodent damage. When bagged
with a high moisture content, seed will heat and soon become worthless.
(Barre, 1954; Wheeler and Hill, 1957; Harrington and Douglas, 1970)
Country Elevator Storage
Many country elevators accumulate seeds during and immediately
after harvest. Storage, which is usually for a limited time, may be
either in bulk, in wood or metal bins, in concrete silos, or in bags or
toteboxes in wood, brick, concrete, or metal buildings. Country eleva-
tors frequently have a fanning mill to remove trash before storing
seeds. Fanning also helps reduce seed moisture content. Country ele-
vators frequently have facilities for insect and rodent control and may
have drying equipment (Barre, 1954).
Seed Processor Storage
Seed processors as a group store a major part of the seeds held for
future use in the United States. Their storage facilities vary widely in
size and construction, ranging from small wood or metal buildings to
large multistoried wood, brick, or concrete warehouses. Some proces-
sors use well-insulated rooms, in which the temperature and relative
humidity are controlled for maximum viability protection, especially for
high-cost items. Seed processors store both processed and unprocessed
seeds. Uncleaned seeds may be put in bags, pallet boxes, or bulk bins.
Each container has certain advantages and disadvantages for specific
kinds of seeds. Uncleaned seeds are usually stored in an area well
separated from where the cleaned seeds are stored. Completely proc-
essed seeds are generally stored in the container in which they will be
shipped. Some processed seeds are stored by the processor in retail
packages, but other seeds are repackaged by the retailer. Storage
facilities operated by seed processors usually have the basic require-
ments previously discussed and may also provide for special needs of
individual kinds of seeds and for protection from unfavorable climatic
conditions.
Retail Storage
Seed retailers must hold their stock for varying lengths of time,
usually under normal atmospheric conditions for their individual loca-
tions. Retail storage is usually far from ideal; however, except in very
hot, humid areas, the shelf life of seeds in cloth, paper, or thin plastic
containers is usually sufficiently long to permit marketing with little loss
of viability. In hot, humid areas, shelf life of seeds is limited unless they
are dried to a safe level of 5- to 8-percent moisture and put in mois-
ture-barrier packages. At the retail level little or no attempt is made to
provide protective storage.
Research Storage
Scientists must store varying quantities of specific seed stocks in
order to have viable seeds available for their research. Although
refrigerated, dehumidified storage is best suited for long-term holding
of seeds, such conditions are not uniformly available to research scien-
tists. They frequently must use less than ideal conditions, although
ordinarily they use the best facilities available.
In some geographic areas, numerous kinds of seeds can be stored
without less of viability for several years under normal atmospheric
conditions. In tropical and subtropical areas, controlled atmospheric
storage conditions are essential to preserve research seed stocks for
periods longer than a few days. The only alternative method is sealed
storage. For safe, sealed storage for up to 3 to 5 years at ambient
temperatures, seeds must be dried to 5- to 8-percent moisture content
before they are sealed in moistureproof containers. For longer storage,
the moisture content must be reduced to 2.5 to 5 percent before
packaging.
Germ Plasm Storage

Germ plasm preservation centers, such as the U.S. National Seed
Storage Laboratory (fig. 24) (James, 1972), the U.S. Regional Plant
Introduction Stations, the Japanese National Seed Storage Laboratory
(Ito, 1972), and similar facilities throughout the world, must have the
best possible storage conditions to minimize regrowing of seed stocks.
Many kinds of :,,eels held under the conditions used in the U.S. National
Seed Storage Laboratory do not have to be regrown more frequently
than every 20 or 30 years.
This laboratory is a Lhree-level building. The machine room (fig. 25),
control room, biochemistry laboratory, growth chamber room, custo-
FIGURE 24 .—U.S. National Seed Storage Laboratory, Fort Collins, Colo.
FIGURE 25. —Machine room of the U.S. National Seed Storage Laboratory, showing some
of the equipment required for a refrigerated storage facility.
dian's supply room, workshop, garage, and a supply storage room are on
the first level. The second level houses the administrative offices. The
seed storage rooms (fig. 26) and germination laboratory (fig. 27) occupy
the third level.
The seed storage rooms, accessible from a common corridor, have a
total capacity of approximately 180,000 pint cans for samples. Seed
sample capacity can be increased manyfold by using containers tailored
to fit the sample. Storage conditions are maintained at about 4° C, with
an average relative humidity of approximately 35 percent. This combi-
nation was selected as suitable for storing most kinds of seeds for a
relatively long time. Three of the rooms are equipped to maintain a
temperature as low as —12° if desired. For research purposes a variety
of temperatures and relative humidities are available in smaller rooms
not used for routine storage of germ plasm.
Constructing Controlled Atmosphere Seed Storage

Facilities
Safe storage of seeds requires careful control of both the temperature
and the relative humidity of the storage area. They cannot be controlled
except in specially constructed rooms or buildings. Because of the need
FIGURE 26. —Sample storage room of the U.S. National Seed Storage Laboratory, where
a scientist is checking samples stored in tin cans.
for effective barriers from outside sources of heat and moisture, the
walls, ceiling, and floor of a seed storage room must have satisfactory
heat insulation and a moisture vapor seal. Figures 28-30 illustrate
construction techniques that will provide the necessary heat and mois-
ture barriers.
Floor insulation is frequently installed in a bed of hot asphalt, which
provides a good vapor seal. The amount of insulation used depends on
the temperature to be maintained and the type of material used, such as
fiber glass, spray on foam, Styrofoam, and cork. Insulating materials
FIGURE 27.—Laboratory workers in the U.S. National Seed Storage Laboratory testing
seeds for germination or decline in viability as storage time is increased.
must be kept dry for maximum efficiency. If the material does not have
a characteristic for dryness built into it, moisture protection must be
provided outside the insulation.
Board-type insulation should be applied in two or more layers, with
the joints lapped or staggered to minimize heat and moisture penetra-
tion through the joints. To cope with the problem of building movement
with changing temperature, an accordianfold is used in the corner
flashing vapor seal material. Ceiling insulation can be of many kinds.
Ceiling and wall finishes usually consist of one-half inch or more of
cement plaster applied as two coats. Where the wall is subject to shock,
the finish coats are reinforced with galvanized metal lath. Wood, metal,
or concrete bumpers are installed on walls where trucks might acciden-
tally hit them.
Cold storage rooms must have no windows and their doors must be
well insulated and well sealed. For large openings, roller hung doors
may be better than swinging doors. Roller doors not only fit tighter but
can be operated electrically. A relatively new idea is the use of a high
velocity stream of cool air across the face of the door, usually from top
to bottom. This may not be the complete solution to the entrance of heat
and moisture, but it does provide some protection. Double-door air locks
FIGURE 28. —Floor construction of a refrigerated seed or grain storehouse, emphasizing

importance of tight seals and insulation. (Courtesy of Amspec, Inc.)
and small anterooms also help reduce heat and moisture entering cold
storage rooms.
The biggest problems in refrigerated dehumidified storage are heat
and moisture leakage through the walls, roof, floor, and around the
doors; heat generated by the lights; and heat and moisture generated by
people working in the room. These, of course, can best be reduced by
incorporating adequate preventative measures into actual construction
of the storage room or warehouse. It is usually desirable to construct
several controlled temperature rooms rather thy;, a single large ware-
house. By having several individually controlled rooms, annual operat-
ing costs can be lowered significantly. During periods when only small
quantities of seeds are stored, one or two rooms rather than an entire
warehouse can be refrigerated.
Most refrigerated seed storage facilities use forced air circulated
through a cooling coil, then throughout the room. For large areas, a
duct system distributes the cold air uniformly throughout the room. The
final decision as to the structural design of the building or room and the
size and type of refrigeration system must be left to a competent
refrigeration engineer. It is far better and more economical to install
initially adequate insulation, moisture protection, and refrigeration
FIGURE 29.—Metal pan ceiling for blast freezer, emphasizing supporting structure, close
fitting seals, and insulation. (Courtesy of Amspec, Inc.)
capacity than to remodel an unsatisfactory system. For further infor-

mation, see May (1964), Dryomatic Division, LogEtronics (1965), Mun-
ford (1965), Cooke (1966), and Harrington and Douglas (1970).
Controlling Temperature
Refrigeration
Since refrigeration in broad terms is any process by which heat is
removed, it is the only way to obtain and maintain the low temperature
required for long-term storage of seeds. Ventilation can be considered
to be refrigeration; however, ventilation is suitable for only minor
downward adjustments of temperature. To obtain very low tempera-
tures, mechanical refrigeration must be used in addition to air circula-
tion. Refrigeration is accomplished by transferring heat from the body
being refrigerated to another body where the temperature is below that
of the body being refrigerated. Because heat moves readily from an
FIGURE 30.—Construction utilizing suspended T-bar ceiling and foam plastic insulation
for seed storage facility. (Courtesy of Amspec, Inc.)
area of high temperature to one of lower temperature, thermal insula-

tion is always necessary around an area to be refrigerated.
Heat Load
The heat load is the rate at which heat must be removed in order to
produce and maintain the desired temperature. Heat load is affected by
such factors as temperature of the space or material to be refrigerated;
heat leakage through the walls, floor, ceiling, and door opening of the
chamber; heat from lights, motors, and other electrical equipment; and
people working inside the refrigerated chamber. Total heat from these
sources determines the heat load for a particular refrigeration system.
Refrigeration Agent
The body employed to absorb heat is the refrigeration agent or
refrigerant. Cooling systems are classified as either sensible or latent,
according to the effect the absorbed heat has on the refrigerant. The
cooling process is said to be sensible when the absorbed heat increases
the temperature of the refrigerant and latent when the physical state of
the refrigerant is changed. With either process the temperature of the
refrigerant must always be lower than that of the area or material being
refrigerated. Continuous refrigeration can be achieved by either a
sensible or a latent cooling system. With a sensible cooling system, the

refrigerant is chilled and recirculated. Latent cooling may be accom-
plished with either a solid refrigerant, such as ice or solid carbon dioxide
(dry ice), or a liquid. Ice and dry ice are not recommended as refriger-
ants for seed storage rooms.
Liquid Refrigerants
Mechanical refrigeration systems are based on the ability of liquids to
absorb enormous quantities of heat as they vaporize. Vaporizing liquids
provide a refrigeration system that can be easily controlled. Such
systems can be started and stopped at will and the rate of cooling can be
predetermined within narrow limits. The vaporizing temperature of the
liquid can be regulated by controlling the pressure at which the liquid
vaporizes. By using a closed system, the vapor can be readily condensed
back into a liquid so that it can be used over and over again to provide a
continuous flow of liquid for vaporization.
Since no one liquid refrigerant is best suited to all applications and
operating conditions, the refrigerant selected should be the one best
suited to meet the specific needs of each storage facility. Of all the fluids
currently used as refrigerants, the one nearest the ideal general pur-
pose refrigerant is dichlorodifluoromethane (CC12F 2). It is one of a
group of refrigerants introduced under the trade name "Freon," but it
is now manufactured under several other proprietary names. To avoid
any confusion among trade names, this compound is now referred to as
Refrigerant-12 or R-12.
Refrigerant-12 has a saturation temperature of -29.8° C, which
means it can be stored as a liquid at ordinary temperatures only under
pressure in heavy steel cylinders. Although an insulated space can be
refrigerated by allowing liquid R-12 to vaporize in a container vented to
the outside, this is not a practical method of refrigerating a seed storage
room. The container in which the refrigerant vaporizes during refrig-
eration is called the evaporator and is an essential component of any
mechanical refrigerating system.
Typical Mechanical Refrigeration System
A typical mechanical refrigeration or vapor-compression system con-
sists of the following essential parts: (1) An evaporator to provide a heat
transfer surface through which heat moves from the space being
refrigerated into the vaporizing refrigerant; (2) a suction line to convey
the refrigerant vapor from the evaporator to the compressor; (3) a
compressor to heat and Compress the vapor; (4) a hot gas or discharge
line to carry the high-temperature, high-pressure vapor from the com-
pressor to a condenser; (5) a condenser to provide a heat transfer
surface through which heat passes from the hot gas to the condensing
medium; (6) a receiving tank to hold the liquid refrigerant for future
use; (7) a liquid line to carry the liquid refrigerant from the receiving
tank to the refrigerant metering device; and (8) a refrigerant metering
device to control the flow of liquid to the evaporator. The typical
vapor-compression system is divided into a low and a high pressure
side. The refrigerant metering device, evaporator, and suction line
constitute the low part of the system; the compressor, discharge line,
condenser, receiving tank, and liquid line constitute the high pressure
side of the system.
Condensing Units
A condensing unit may be either air or water cooled. Condensing
units of small horsepower are frequently equipped with hermetically
sealed motor compressor assemblies. Large condensing units are
usually water cooled.
System Capacity
The capacity of a refrigeration system is the rate at which it removes
heat from the refrigerated space. Capacity is usually expressed in
terms of units of heat removed per hour or its ice-melting equivalent. In
other words, a mechanical refrigeration system that will cool at a rate
equivalent to the melting of 1 ton of ice in 24 hours is said to have a
capacity of 1 ton. The capacity of a mechanical refrigeration system
depends on the weight of refrigerant circulated per unit of time and
refrigerating effect of each pound circulated.
Compressor Capacity
The capacity of a compressor must be such that the vapor is drawn
from the evaporator at the same rate at which it is produced. If the
vapor is produced faster than the compressor can remove it, the
accumulation of excess vapor will increase the pressure in the evapora-
tor; this will, in turn, increase the boiling temperature of the refriger-
ant. If the capacity of the compressor is such that the vapor is removed
too rapidly from the evaporator, the pressure in the evaporator will
decrease and lower the boiling temperature of the refrigerant. In either
case, the refrigeration systems will not function properly. For any good
refrigeration system, the rate of vaporization must balance the rate of
condensation of the vapor back to a liquid. Such a balanced system will
function properly and refrigerate exceptionally well. For further infor-
mation, see Dossat (1961).
Controlling Humidity
Air is a mixture of gases, consisting primarily of nitrogen, oxygen,
carbon dioxide, water vapor, and small percentages of rare gases. Each
gas, including water vapor, exerts its own partial pressure in the
mixture just as though the other gases were not present. The sum of
these partial pressures equals the total pressure of the mixture. The
amount of water vapor that can be contained in the air mixture is a
constant value depending only on the temperature and pressure of the
mixture. Thinking of humidity in terms of partial pressure makes it
easier to understand the movement of moisture from one area to
another, for moisture moves from a high to a low pressure area. It is
therefore possible for moisture to move against the flow of air. Pres-
surizing a room will not prevent moisture from moving in, although it
could slow down its entry.
Relative humidity is normally measured by taking dry-bulb and
wet-bulb temperature readings and finding the intersection of those
readings as plotted on a psychrometric chart. The point of intersection
will correspond to a particular relative humidity. Relative humidity can
be changed by raising the air temperature without changing absolute
humidity. For further information, see Dryomatic Division, Log-
Etronics (1965).
Moisture Movement Between Air and Materials

Structural materials, such as wood and cement, contain moisture
throughout, whereas such materials as steel and glass hold moisture on
their surface. The rate of moisture vapor movement between such
materials and air is determined by the difference in moisture vapor
pressure between them. If their moisture vapor pressures are equal,
moisture will not move from one to the other.
When moist seeds are placed in a dry atmosphere, moisture will flow
from the seeds into the atmosphere. Because the air cannot hold nearly
all the moisture held in the seeds, the air will soon become saturated
with the moisture given off by the seeds, and unless new dry air is
provided, drying of the seeds will stop. Because construction materials
contain moisture, that Moisture as well as the moisture in the seeds has
to be removed. Once the room and the seeds reach moisture equilibrium
with the desired relative humidity, the drying system has only to
remove the moisture that enters the controlled atmosphere room
through door openings, leakage through seams and cracks, and pene-
tration of the barrier material. The storage area is kept at a designated
relative humidity, which, in turn, prevents any change in the moisture
content of :;toyed seeds once they have reached moisture equilibrium
with the maintained relative humidity. For additional information, see
Hass (1961, 1965), Sijbring (1963), Dryomatic Division, LogEtronics
(1965), and Munford (1965).
Refrigeration-Type Humidity Control Systems

The refrigeration-type dehumidifier draws warm, moist air over a
metal coil with fins spaced far enough apart to permit partial frosting
and still allow for sufficient passage of air. The frost can be removed by
electrical heater, hot gas, or water defrosting at regular intervals
regulated by a timeclock. To be effective at low temperatures, a
refrigeration-type dehumidification system must cool the air below the
desired temperature and reheat it to the desired temperature. Air-
handling units are available with built-in refrigeration coils, electric
defrosters, and reheat coils. An engineered unit such as described may
be better than a piecemeal assembly because the correct wattage is
provided in the unit.
The simplest method of regulating the reheat in a room is to wire the
reheat coils through a relay to a humidistat. Although the refrigera-
tion-reheat system can effectively control relative humidity as well as
temperature, it is not the only system available and in some cases may
not be the best. When very low relative humidity is to be maintained,
operation of the refrigeration-reheat system becomes very expensive.
Desiccant-Type Humidity Control Systems
Dehumidifiers using liquid or solid desiccants in conjunction with
refrigeration can frequently reduce the cost of maintaining very low
relative humidities. These desiccants absorb moisture vapor from the
airstream and later eject it outside the room. Desiccant dehumidifiers
generally use dry chemicals for small systems and salt solutions for
systems with extremely large volumes of air. For seed storage facili-
ties, dry desiccant systems are almost invariably used. The dehumidi-
fier incorporates one or two beds of granulated silica gel or activated
alumina, which can absorb much water vapor. For example, silica gel
can absorb as much as 40 percent of its weight in water vapor at
100-percent relative humidity and proportionally less at lower relative
humidity.
With the two-bed system most frequently used for seed storage
facilities, the air is circulated through one bed at a time. One bed is
recharged, or dried out, while the other is taking up moisture. The
switch from one bed to the other is usually programed by a timeclock for
maximum efficiency.
Recent developments receiving wide application are the rotary drum
or cylinder (fig. 31) and rotary disc (fig. 32) dehumidifiers. The rotary
dehumidifiers have one or more beds divided into two airstreams by
sealing strips. The bed or beds rotate slowly, and while part of each bed
is absorbing water vapor from the airstream, the remainder is being
recharged. The end result is much the same as for the two separate bed
systems except that the rotary systems seem to have a higher drying
FIGURE 31.—Above, rotary cylinder dehumidifier used to control relative humidity in

seed storage rooms; below, arrangement of cylinder, reactivation heaters, and seals
in this dehumidifier. (Courtesy of Dryomatic Div.)
capacity for a given volume of air passing through the dehumidifier. In

contrast to the refrigeration type, desiccant dehumidifers are not
affected by the freezing of moisture and can handle air at —40° to +80°
C as long as the appropriate desiccant is used.
FIGURE 32.—Above, rotary disc dehumidifier used to control relative humidity in seed
storage rooms; below, arrangement of discs and seals in this dehumidifier. (Courtesy
of Dryomatic Div.)
For seed storage facilities where both a low temperature and a low
relative humidity are needed, a combination refrigeration-desiccant
dehumidifier system will probably provide the most reliable tempera-
ture and relative humidity control at the least cost. Maintenance cost of
dehumidification systems is usually small. For desiccant systems, the
desiccant material should be changed every 3 to 5 years. For additional
information, see Hass (1961, 1965), James (1962), Cooke (1966), Beck
(1966, 1972), and Welch (1967).
Common-Sense Practices
Ventilation
Seeds stored wet tend to get hot whether stored in bulk or in bags
unless heat is rapidly dissipated as it is generated. Both heat and excess
moisture can be dissipated through ventilation. A steady stream of air
moving through a bin of loose seeds or a warehouse full of bags of seeds
will collect the heat as it is generated and transport it away. Preventing
heat buildup is essential to maintaining good germination. A good
ventilating system will soon pay for itself through improved seed
quality.
Stacking Seed Bags
Most processed seeds are handled in bags, which are stored in stacks.
Bags of seeds of each genus, species, and cultivar are stored in separate
stacks. To allow for proper ventilation, stacks of seed bags should be
well spaced, both between the bags and between the stacks. Bags of
seeds must be stacked carefully to prevent slippage and falling. Falling
bags are hazardous to employees as well as subject to breakage on
impact. Stacks that are excessively high often cause the bottom bags to
burst. It is especially important that bags moved by forklift be stacked
securely on pallets.
Removing Seeds From Controlled Storage
Dry seeds removed from cold storage and exposed to a warm, humid
atmosphere will absorb moisture readily. Unless preventive steps are
taken, bags of cold seeds will become moist from the condensing
atmospheric moisture. Moisture condensation can be prevented by
warming the seeds in a dry atmosphere or by warming in a room
ventilated with rapidly moving air. Use of moisture-barrier containers
can prevent condensed moisture from coming into contact with the
seeds. Proper aeration can bring cold bulk seeds up to normal atmos-
pheric temperature without surface accumulation of condensed atmos-
pheric moisture. Unless seeds held under controlled atmosphere
storage are kept dry after removal from storage, the protective effects
of controlled storage will soon be lost.
Warehouse Cleanliness
Warehouse cleanliness, or lack of it, can markedly affect seed quality.
A clean warehouse has fewer rodents and insects and they are more
easily controlled. A clean warehouse reduces the chances of accidental
mixtures and personnel accidents. It also presents a better company
image to visitors.
PACKAGING AND PACKAGING MATERIALS

Requirements of Different Situations
Seed-packaging methods are many and varied (Bass et al., 1961). The
equipment for filling packages ranges from a simple spoon or seed
scoop, to the manually controlled gravity flow from a bin, to the
high-speed automatically controlled small packet filler. Package-filling
equipment has no effect on the genetic or physiological quality of seeds;
however, it may affect the physical quality of seeds through impact or
undue pressure. Heavy seeds, especially bean, corn, pea, and soybean,
can be fractured by impact on the individual seed by either striking or
being struck by a hard object or firm surface. Seeds containing too much
or too little moisture damage easily on impact. Damage from repeated
impact is accumulative. Seeds may also be injured by force feeding
through a restricted opening or between pressure rolls. Physical dam-
age can occur during any handling operation from harvest to planting.
Types of Packages
Packages for processed seeds may be burlap, cotton, paper, or film
(plastic, foil) bags, metal or fiberboard cans or drums, glass jars,
fiberboard boxes, or containers made of various combinations of mate-
rials. The types and sizes of containers used for wholesale distribution
of field seeds are usually the same as those used for retail sales. For
vegetable and flower seeds, wholesale containers may be large fabric or
multiwall paper bags, large fiberboard or metal cans or drums, or
fiberboard boxes, whereas retail containers are usually small paper,
fil m, or film-laminated envelopes, small fiberboard boxes, or small metal
cans.
The packaging materials, methods, and equipment used are dictated
by the kinds and amounts of seeds to be packaged, the type of package,
duration of storage, storage temperature, relative humidity of the
storage area, whether packaging is for wholesale, retail, or local use,
and geographical area where the packaged seeds will be stored, exhib-
ited, or sold. Packages that will contain seeds and protect most
physical qualities of seed lots are made of materials with sufficient
tensile strength, bursting strength, and tearing resistance to withstand
normal handling. However, such materials do not protect seeds against
either insects and rodents or changes in moisture content unless special

protective qualities are built into them. Researchers frequently use
whatever container happens to be convenient without regard to the
moisture protection it may or may not afford. However, there is an
increasing awareness of the savings in time and expense that are
realized by using suitable moisture-barrier containers for storing valu-
able breeding stocks.
Seeds stored for a short time or under cold, dry conditions will retain
good viability in porous paper or fabric containers, whereas seeds
stored or marketed under tropical conditions (Ching, 1959) will lose
viability rapidly without maximum moisture protection. Except in
tropical and subtropical climates, many kinds of seeds do not require
special moisture protection during the first winter after harvest when
held in the area where produced. However, seeds carried over to the
second planting season often require drying and packaging in mois-
ture-barrier containers to prevent loss of viability and vigor.
How Packages Are Filled
Except in very limited operations that employ hand-filling, seeds to
be packaged are delivered to hopper bins above automatic or semiauto-
matic filling machines. Seeds may come to the hopper from bulk storage
bins by gravity flow through pipes and by airlift, belt conveyors,
forklifts, or the human shoulder. Since practically all seeds are sold by
weight, it is necessary to put a predetermined amount into the individ-
ual packages. Even seeds that appear to be sold by volume are asso-
ciated with weight; namely, a bushel of corn equals 56 pounds, a bushel
of wrinkled peas 56 pounds, and a bushel of smooth seeded peas 60
pounds. Most package-filling equipment has built into it a seed-measur-
ing device, or it is manually or automatically controlled by a signal from
a weighing device.
How Containers Are Presented to the Filler
Large nonrigid containers of burlap, cotton, lined bags, multiwall and
7- and 10-mil polyethylene bags are usually presented to the filler
manually. They are held in place by hooks, clamps, or by hand during
filling. Polyethylene and similar materials may be formed into bags
from sheets or rolls, filled, and sealed in a continuous operation. Large
rigid containers, other than metal or glass, are usually formed into an
open box by hand and presented to the filling equipment either manually
or by a conveyor that automatically positions each container. Small,
semirigid, preformed containers may be opened with a jet of air and
automatically positioned for, filling. Rigid containers come from the
manufacturer ready for filling. They are positioned for filling either
manually or by conveyor.
Placing seeds in rolls of tape may be considered a form of packaging.
During the midtwenties, equipment was developed for forming the

tape, placing the seeds, and making the roll. Although this form of
packaging did not. become popular at that time, there currently appears
to be a revival of interest in this method.
Weighing and Measuring Devices
Scales for weighing seeds range from the large truck scale, which
weighs an entire load at a time, to the common beam scale or an
elaborate scale that activates either a pneumatic or electric device that
shuts off the flow of seeds when a predetermined weight or volume is
reached. In retail stores many types of scales are used, some of which
automatically compute the cost to the customer based on the price per
pound. Some scales are so delicate they can weigh a few small seeds,
such as petunia. Measuring devices range from a simple spoon or scoop,
to a calibrated cylinder or cup, to the measuring device associated with
the automatic scales.
How Containers Are Closed
Hand-tying of cotton and burlap bags has been largely replaced by
sewing. Although there is still some sewing by hand, most of it is done
by machine. Polyethylene and other thermoplastic materials are usually
sealed by applying heat at 93.3° to 204.4° C to the film for a given time
while the film is under pressure. Within limits each kind and thickness
of material has specific temperature, time, and pressure requirements
(table 23) for proper sealing.
TABLE 23. —Approximate heat-sealing requirements for selected

thermoplastic materials1
Heat sealers range from small hand-operated rollers or bars, to

foot-operated bars, jaws, and clamps, to elaborate automatic bag- or
pouch-forming, filling, and sealing machines. Some sealers use ther-
mostatically controlled bars, bands, or rollers, and others use high-in-
tensity thermal impulses of short duration. Most are readily adjustable
for use with many kinds and thicknesses of material. However, the
quality of the seal produced by hand- and foot-operated equipment
depends on the skill of the operator. Properly adjusted automatic
sealers seldom produce a poor seal. Considerable experience is required
before an operator can consistently apply the appropriate pressure for
the precise dwell time required to produce a good seal. It is especially
important that seed packages of proper moisture-barrier materials be
well sealed as a leak in the seal will soon negate the moisture protection
provided by the material.
Semirigid or rigid containers, other than metal or glass, are usually
sealed with cold or hot glue applied either by hand or automatically by
machine. Most machines that apply glue also form and fold the open
ends of the containers and place them under pressure until the glue has
set. Rigid containers, such as fiber drums, may have slip-on caps or lids
that clamp into position. These lids are applied manually, whereas lids
for metal cans are applied mechanically. Can sealers are manually
operated, semiautomatic, or fully automatic. Usually the equipment is
the same as that used by food processors for hermetically sealed cans. A
partial vacuum or a gas can be introduced into the can when semiauto-
matic and fully automatic equipment is used for sealing. Some cans are
sealed with pressure lids similar to those used on paint cans.
Packaging Field Seeds
Field seeds are generally packaged in burlap, osnaburg, or seamless
and multiwall paper bags containing 50 or 100 pounds or one-half to 3
bushels of seeds. A few companies use moisture-barrier packages, such
as elastic multiwall paper bags with either an asphalt or a polyethylene
ply in the multiwall, burlap or cotton bags with polyethylene liners, and
burlap/asphalt/paper-laminated bags for cereal, soybean, and hybrid
sorghum seeds. Many hybrid corn seed producers either package all or
part of their crop each year in moisture-resistant packages or hold their
seed in controlled storage. Moisture-resistant bags used for seed corn
are made of multiwall/asphalt/paper, elastic multiwall/asphalt/paper,
multiwall and elastic multiwall/paper with a polyethylene-barrier ply,
or 7- or 10-mil polyethylene.
A valve-type polyethylene bag has been developed that prevents any
loss of material while filling and that seals easier than the conventional
bag. The valve-type bag also permits easy introduction of fumigants or
gases into the filled bag prior to sealing.
Some companies are packaging hybrid corn in acre units, each pack-
age containing the exact quantity of seed required to plant a specified
number of acres.
Turf grass seed for the wholesale market is usually put in fabric bags
of 25- to 100-pound capacity. Sometimes a 4- or 5-mil polyethylene liner
is used for moisture protection. At retail, turf grasses are sold as single
species or combinations of species packaged in paper, cloth, or polyeth-
ylene bags, metal cans, or cardboard boxes. The cardboard boxes may
be plain, polyethylene, or wax paper lined, foil or wax coated, or wax
paper or foil overwrapped.
Tobacco seeds are packaged in paper packets, cardboard boxes or
cylinders, or metal cans of 1/2- or 1-ounce capacity.
Moisture-resistant containers, such as wax- or polyethylene-coated
cardboard boxes or drums and polyethylene, cotton, or paper combina-
tions, are being used for various forage grasses and legumes. Cotton-
seeds previously dried to 6- to 8-percent moisture content are packaged
in burlap/asphalt/paper bags or wax- or polyethylene-coated boxes.
Large wood or steel pallet boxes with covers are being used extensively
in the seed industry for bulk storage of both uncleaned and cleaned
seeds. Such bulk storage reduces handling costs and increases the
efficiency of cleaning plant operations by permitting a greater amount of
mechanization. There is no handwork involved in handling pallet boxes,
but with bags or small boxes, a great deal of handwork is required.
Packaging Vegetable and Flower Seeds
In the vegetable seed industry, the packaging method is determined
primarily by the type of customer for which the seeds are packaged.
Seeds for wholesale distribution are usually packaged in fabric bags of
25- to 100-pound capacity. The bags may or may not have polyethylene
liners. Beans, peas, and sweet corn are usually packaged in mul-
tiwall/asphalt/paper bags containing 25 to 100 pounds of seed. For retail
sales, most vegetable seeds are packaged in paper packets, sometimes
with foil inserts, containing a fraction of an ounce to several ounces;
cardboard boxes holding a few ounces; and 2- to 4-mil polyethylene,
cellophane, acetate, paper, and foil-laminated bags with capacities of
to 5 pounds. Various kinds of vegetable seeds are also packaged in
hermetically sealed metal cans of various sizes. These containers are
especially beneficial for shipments overseas and into tropical areas.
Flower seeds at the wholesale level are packaged in paint-type cans,
glass jars, and fabric bags with polyethylene liners. For retail sales,
paper, paper/polyethylene, acetate, cellophane, and foil-laminated
packets are the predominate types. Wholesale packages of flower seeds
usually contain several pounds, whereas retail packets contain a given
number of seeds or a fraction of an ounce.
Package Labeling
Researchers and seedsmen alike need to identify the contents of each
container, both as to species and cultivar. Frequently other informa-
tion, such as percentage of live seeds, purity, noxious weed seed
content, and seed treatment, if any, must also be recorded as required
by law. This may be done by printing the required information on a tag
attached to flexible cotton or fiber bags, by printing on a label glued to
tin cans, cardboard boxes, or cardboard or metal drums, or by printing
or stamping the information directly on the container, such as litho-
graphing cans or embossing metal lids.
Porous Packaging Materials

Burlap or hessian bags are made of good quality jute yarn in a variety
of fabric constructions. Since burlap is exceptionally strong, burlap
bags lend themselves to stacking high in storage and to rough handling,
and they can be reused many times. Seed bags of cotton are made of
sheeting, printcloth, drill, and osnaburg fabrics as well as a special
seamless material. Osnaburg and seamless fabrics have the greatest
strength and tear resistance of the cotton materials. Bags made of these
materials are reused many times, whereas bags of other cotton materi-
als are used only once. Reuse of fabric bags is largely confined to
storage of unprocessed seed as certification standards require new bags
for shipment of processed seeds.
Paper products are extensively used for seed packaging. Small seed
packets are mostly made of bleached sulfite or bleached kraft paper and
surface coated with a very white clay to facilitate printing. Basically
paper packets are designed to contain, without loss, a given quantity of
seeds, not to protect sued viability.
Many paper seed bags are of Tnultiwall construction consisting of
several layers of smooth or crinkled paper. Multiwall paper bags are
produced in a variety of constructions, each designed for a specific
purpose. Ordinary multiwall bags have poor bursting strength and
consequently when piled high, the bottom bags burst. The top bags in
high piles often slip. Under very dry conditions, multiwall bags dry out
and become brittle along folds and at wear points. Elastic multiwall
bags consist of several layers of crinkled paper. Elastic materials cannot
be evaluated by the usual physical test data for tensile and tear strength
because the entire principle of the elastic multiwall bag depends on
built-in stretchability.
Cardboard, in the form of boxes and cans, is used extensively in seed
packaging. Cardboard containers protect most physical qualities of seed
and are well adapted for automatic filling and sealing equipment. Porous
packaging materials adequately contain or hold the seeds and protect
them from mechanical mixtures, but they do not provide moisture

protection.
Moistureproof Materials
Metal containers when properly sealed provide an absolute moisture
and gas barrier and completely shield the products from any effects of
light. Metal containers provide complete protection against rodents,
insects, changing humidity, flood, and harmful fumes. Metal cans are
well adapted to high-speed automatic filling and sealing.
Glass containers are not widely used in seed packaging. Although
glass provides essentially the same protection as metal, its fragility
makes it less desirable for commercial packaging. Glass containers are
frequently used in research and as display receptacles in seed and
hardware stores, where bulk hales of seed are made. Glass containers
are often used by home gardeners for carryover of small quantities of
seed from one season to the next.
Adequate drying before sealing is absolutely essential for safe
storage of seeds in airtight, moistureproof containers, especially for
seed stored at warm temperatures or shipped to tropical areas. Much
research has been done on the effects of sealed moistureproof storage
on seed longevity. Some workers used air-dried seeds, whereas others
carefully dried the seeds before sealing.
Moistureproof Storage
Response of Different Crop Seeds to Moistureproof Storage
Cereal Seeds.—Numerous studies have shown that rice seeds held in
airtight storage outlive seeds held in open storage provided they are
adequately dried before they are placed in airtight storage (Vibar and
Rodrigo, 1929; Kondo and Okamura, 1930, 1932-33, 1938; Rodrigo,
1935, 1953; Kondo and Terasaka, 1936). Rice seeds dried by high heat
before sealing were dead after 26 to 28 years, but seeds dried by the sun
to 11- to 13-percent moisture were stored safely for 30 years (Kondo and
Okamura, 1932-33). Cultivar differences in storability were demon-
strated by Vibar and Rodrigo (1929), who reported that the germination
of seeds of the rice cultivar Hambas declined 18 percent in 51 months,
whereas seeds of the Inintew cultivar declined only 5 percent.
Because corn seed is usually planted the year after harvest, people in
the Corn Belt usually have no problem with storage. However, corn is
now grown in areas where temperature and humidity conditions fre-
quently cause rapid loss of viability during storage. Research has shown
that sealed storage prevents rapid loss of viability of corn seeds
provided their moisture content is sufficiently low when the seeds are
sealed (Vibar and Rodrigo, 1929; Rodrigo, 1935, 1953; Kaihara, 1951;
Barton, 1960b). Sealed seeds with 11-percent moisture content main-
tained full viability for 9 years at –5° C but showed reduced viability
after 1 year at 30° (Barton, 1960b). Although sealed glass jars were
used in this study, similar results could be expected with any type of
sealed moistureproof container, such as a metal can or drum or a
multiwall bag with a foil layer.
Sorghum seeds in commerce usually are not stored longer than over
winter. However, plant breeders frequently need to hold seeds for
many years. Sorghum seeds dried and sealed in glass bottles retained
their viability longer than did similar seeds stored in gunny sacks
(Krishnaswamy, 1952). We found that cultivar RS619 sorghum seeds
retained their germination up to 8 years at –12° and –1° C, whether in
paper envelopes or sealed metal cans (table 24). Sealed seeds at 10°
retained viability significantly better than did unsealed seeds. At higher
temperatures the results were variable. Some of the variability can be
attributed to differences in seed moisture content. The seeds in sealed
metal cans had an initial moisture content that did not change with time.
The moisture content of the seeds in paper envelopes adjusted with
time to moisture equilibrium with the relative humidity in the storage
chamber. he low relative humidity at the higher temperatures allowed
the seeds to attain a low moisture content and partly offset the effects of
temperature on longevity.
TABLE 24 . —Germination of 5 kinds of seeds at 3 initial moisture

contents after storage in paper envelopes or sealed metal cans
under 5 temperature and relative humidity conditions for 4 and 8
years
TABLE 24. —Germination of 5 kinds of seeds at 3 initial moisture

years—Continued
TABLE 24.—Germination of 5 kinds of seeds at 3 initial moisture

years—Continued
TABLE 24. — Germination of 5 kinds of seeds at 3 initial moisture

years—Continued
Forage Grasses.—Seeds of Echinochloa, Eleusine, Panicum, Pen-

nisetum, Paspalum, and Setaria retained their viability longer when
dried and stored in sealed bottles than when stored in gunny sacks.
Sealed Echinochloa, Panicum, and Setaria seeds retained approxi-
mately 70-percent germination for 38 months (Krishnaswamy, 1952).
Crested wheatgrass, intermediate wheatgrass, and smooth bromegrass
seeds retained high viability best at –18° C whether sealed or open
(Knowles, 1967). At 1°, seeds stored in sealed glass jars held their
germination better than those in open storage. At 21°, seeds in sealed
containers retained a higher percent viability than did seeds in paper
envelopes; however, the viability of sealed seeds sharply declined
during 4 years of storage.
Turf Grasses.—Perennial ryegrass seeds containing 6-, 8-, 12-, 16-,

and 20-percent moisture were stored in sealed metal cans at 3°, 22°, and
38° C in a warehouse in Oregon (Ching et al., 1959). After 3 years of
storage, the 6-percent moisture seeds showed some decline in germina-
tion at 38°, seeds with 8-percent moisture retained good germination at
all temperatures except 38°, but seeds with 12- and 16-percent moisture
remained viable only at 3°. Seeds with 16- and 20-percent moisture
deteriorated within 3 months when stored at 22° and 38°. Ching and
Calhoun (1968) reported that after 10 years of sealed storage, 6-pereent
moisture ryegrass seeds stored at 22° and 38° gave significantly lower
laboratory germination percentages than new seeds and similar seeds
held at other temperatures; however, field emergence was not signifi-
cantly different from that of all other seed lots.
Kentucky bluegrass seeds with 8.7-, 6.2-, and 4.9-percent moisture
packaged in sealed tin cans and stored at r, 10 6 , and 226 C germinated
at approximately their original level after 30 months in storage regard-
less of the seed moisture level (Bass, 1960). Creeping red fescue
containing 11.4-percent moisture in sealed metal cans dropped sharply
in viability during the first 3 months of storage at all temperatures;
seeds sealed with lower moisture levels retained good viability longer,
especially at the lower temperatures.
Forage Legumes.—Crimson clover seeds containing 6-, 8-, 12-, 16-,
and 20-percent moisture were stored in sealed metal cans at 3°, 22°, and
38° C in a warehouse. Seeds with 16- and 20-percent moisture at 22° and
38° deteriorated within 3 months, those with 8-percent moisture were
preserved well except at 38°, and those with 12- and 16-percent mois-
ture remained viable only at 3°. The 6-percent moisture crimson clover
seeds remained highly viable after 3 years of storage under all temper-
ature conditions. (Ching et al., 1959) Ching and Calhoun (1968) re-
ported that after 10 years of sealed storage only 16-percent moisture
crimson clover seeds held at 3° germinated significantly lower than
fresh seeds in laboratory tests. In the field there were no significant
differences in emergence between fresh and 10-year-old seeds regard-
less of the temperature under which the sealed seeds had been stored.
In our studies, crimson clover seeds with 4- and 7-percent moisture
retained higher viability or germination in sealed metal cans than in
paper envelopes at all storage conditions (table 24). However, the
10-percent moisture seeds lost viability more rapidly in the cans than in
the paper envelopes when stored at 21° and 32° C. This was, of course,
because the seeds in paper envelopes lost moisture during the early
weeks of storage, whereas those in the cans did not. The 10-percent
moisture seeds in metal cans actually declined 50 percent in viability
during the first year of storage at 32°, whereas the seeds in paper
envelopes declined only 11 percent.
These data show that seeds in hermetically sealed containers may lose
viability more rapidly than those in open storage if seed moisture
content, storage temperature, or both is too high.
Alfalfa seeds with 6.8-percent moisture germinated 78 percent after
24 years in sealed glass bottles, and red clover seeds with 6.9- and
6.7-percent moisture germinated 74 and 71 percent, respectively (Nu-
tile, 1958).
Fiber and Oil Crops.—Flaxseeds containing 7- to 8-percent moisture
when stored in metal containers retained good germination for 13 to 16
rears at temperatures prevailing in Mandan, N. Dak. (Dillman and
oole, 1937). Cottonseeds with 6- to 8-percent moisture in sealed glass
jars kept for 7 to 10 years at 21° C without loss of viability (Simpson,
1942, 1953). Good quality cottonseeds with 7- to 11-percent moisture
content retained high viability for over 25 years in sealed metal cans at
1° (Pate and Duncan, 1964).
Hempseeds with 8.6-percent moisture or less retained good germi-
nation for 31/2 years at 2° C (Crocioni, 1950). Seeds with 5.7-percent
moisture still retained their initial viability after 15 years of sealed
storage at –10° and 10° (Toole et al., 1960; Clark et al., 1963). Two lots
of seeds with 6.2-percent moisture in sealed cans retained full viability
for 12 years at –10°, 0°, and 10°, whereas a third lot of lower initial
germination decreased 23 percent in germination during 12 years of
storage. The two high germinating lots of hempseeds when stored with
9.5-percent moisture retained good viability at –10° and 0° but de-
teriorated rapidly at 10°. Seeds of the poorer germinating lot declined
10 percent at –10°, 27 percent at 0°, and lost all viability at 10° during 12
years of storage.
Kenaf seeds with 8-percent moisture retained their initial viability for
12 years when stored sealed at –10°, 0°, and 10° C. Seeds with
12-percent moisture, which had retained full viability for 51/2 years at
–10° and 0° and showed a significant loss of viability after 41/2 years at
10° (Toole et al., 1960), were nonviable after 12 years at 10° and had
declined sharply at 0° (Clark et al.; 1963).
For both safflower and sesame seeds, careful control of seed moisture
is absolutely essential for sealed storage. Safflower seeds containing 4-
and 7-percent moisture retained their germination about equally well in
paper envelopes and in sealed metal cans when stored for 8 years at
–12°, –1°, and 10° C (table 24). The 4-percent moisture seeds in sealed
cans retained better viability than did the envelope-stored seeds at 21°
and 32°. However, 7-percent moisture seeds in sealed metal cans lost all
viability in less than 4 years at 21° and in less than 1 year at 32°.
Ten-percent moisture seeds in sealed metal cans retained fair viability
for 8 years at –12° and –1°; however, all were dead in less than 3 years
at 10° and in less than 1 year at 21° and 32°. The seeds in paper
envelopes at 21° and 32° markedly decreased in viability, but no sample

was completely dead regardless of the initial moisture content or
storage temperature. For safe, sealed storage of safflower seeds at a
temperature greater than 10°, seed moisture content must be reduced
to 4 percent or less before sealing.
Sesame seeds in sealed metal cans did not retain satisfactory germi-
nation when seed moisture content exceeded 7 percent for storage at
–12° and –1° C (table 24). At 10° and higher, seed moisture content
must not exceed 4 percent. Ten-percent moisture seeds in sealed metal
cans do not store well at any temperature. The 10-percent moisture
seeds at 10° germinated 73 percent after 1 year but were nearly all dead
at the end of the second year. Seeds at 21° and 32° lost all viability in less
than a year. As with safflower, sesame seeds are very sensitive to both
seed moisture content and storage temperature and must be carefully
dried to 4-percent moisture before sealing unless the seeds are to be
held continuously at –12° or –1°.
Under Philippine conditions the germination of air-dry soybean seeds
stored in sealed containers lost viability after 23 months. (Vibar and
Rodrigo, 1929) and 54 months (Rodrigo, 1953). In Illinois the germina-
tion of air-dry soybeans in sealed containers dropped only 2 percent
during the first year but declined sharply thereafter, and nearly all
seeds were dead within 8 years (Burlison et al., 1940). The germination
of 'Illsoy' at 7 years was comparable to 'Manchu' at 4 years and
'Lexington' at 5 years, indicating cultivar differences in keeping quality
(Burlison et al., 1940).
Soybean seeds conditioned to various moisture contents, then sealed
in pint jars, and stored at various temperatures lost viability at a
different rate at each storage condition (Toole and Toole, 1946). Cul-
tivars of Mammoth Yellow and Otootan gave essentially the same
results. Seeds with 18-percent moisture at 30° C were nonviable in less
than 3 months, and similar seeds at 10° died within 2 years. Seeds stored
at subfreezing temperatures retained good germination for 6 years but
were unsatisfactory for planting after 10 years. Air-dry seeds of
13.5-percent moisture, however, retained essentially their full initial
viability for 10 years at 2° and –10°. Soybean seeds dried to 9-percent
moisture or less before sealing retained full viability for 10 years at 10°,
2°, and –10° (Toole and Toole, 1946).
Peanuts in sealed storage retained satisfactory germination up to 37
months (Vibar and Rodrigo, 1929; Rodrigo, 1953).
Vegetable Seeds.—Onion seeds kept very well in sealed storage
(Brison, 1941, 1942). Seeds dried to 6.4-percent moisture before sealing
germinated 90 percent after 13 years at room temperature (Brown,
1939). However, in another study (Asgrow Seed Co., 1954), 6.3-percent
moisture content seeds retained good viability for only 3 years at 32° C.
Although onion seeds dried to below 8-percent moisture before sealing

kept well for 4 years at 5° (Beattie and Boswell, 1939), 6.7-percent
moisture is recommended for longer storage. Dry seeds in sealed
containers in a laboratory lost viability in 38 months (Rodrigo, 1953).
Sealed onion seeds held in cold storage at 5° to 10° without deterioration
for 2 years did not deteriorate when held 3 months at natural conditions
after removal from storage (Myers, 1942). This indicates that ade-
quately dried onion seeds held sealed in cold storage can be safely
marketed after removal from cold storage.
Rodrigo (1935, 1953) reported that under Philippine conditions the
viability of air-dried mung beans stored in sealed containers in a
laboratory was maintained up to 201 months. Cowpeas similarly stored
remained viable for 123 months. White-seeded tapilan beans lost via-
bility gradually over 10 years, whereas black-seeded tapilan beans
maintained their original viability (Vibar and Rodrigo, 1929; Rodrigo,
1935). Yellow tapilan seeds lost viability in 138 months, whereas black
tapilan seeds maintained viability up to 201 months (Rodrigo, 1953).
Seeds of 'Buff Cross,' `Mahogany Brown,' and Red Kidney'
beans germinated 20 percent, whereas 'Improved Michigan Robust'
seeds germinated 50 percent after 38 years of storage in sealed canning
jars in a laboratory at Geneva, N.Y. (Waters, 1962). For 'Top Notch
Golden Wax' and 'Bountiful' bean seeds stored open and sealed at —18°,
—2°, 5°, 10°, 20°, and 30° C and ambient temperature in a laboratory at
Yonkers, N.Y., sealing extended longevity when storage was in a
humid room, even as low as 5° (Barton, 1966a). Sealing was without
effect at —18° and —2° for up to 15 years. Storage at 30° caused rapid
deterioration in both open and sealed containers.
The germination and vigor of canned cucumber seeds did not change
significantly in 36 months when seed m oisture was 5.3 percent or less at
32° C or lower (Asgrow Seed Co., 1954). With higher seed moistures,
temperature becomes more critical. Seeds with 9.4-percent moisture
became worthless in 3 months at 32° but held viability and vigor well for
30 months at 15.6°. Tin cans were superior to chlorinated rubber, waxed
cellophane, and paper bags inside linen ones (Coleman and Peel, 1952).
Dry seeds in sealed containers in a laboratory lost viability in 38 months
(Rodrigo, 1953).
For safe sealed storage at room temperature, parsnip seeds ma' be
dried to less than 1.7-percent moisture before being sealed (Joseph,
1929). For safe storage at 5° to 7° C, the critical moisture level was 6.13
percent. When ovendry, 4-, 6-, and 8-percent moisture content parsnip
seeds were stored for 6 years in sealed and unsealed containers at room
temperature, 5°, and 6.7°, the best keeping was obtained with 4-percent
moisture seeds at 6.7°, followed by ovendry at 5° (Beattie and Tatman,
1950). Sealed storage also was effective in prolonging the life of carrot
seeds (Coleman and Peel, 1952; Dutt and Thakurta, 1956). In open
storage viability was lost in 9 months, but in sealed storage the seeds
retained their viability.
Sealed storage prolonged the life of seeds of brussels sprouts, cab-
bage, cauliflower, knolkol or kohlrabi, radish, and rutabaga or swede
(Coleman and Peel, 1952; Rodrigo, 1953; Dutt and Thakurta, 1956;
Madsen, 1957). Loss of viability of dry seeds in sealed containers in a
laboratory occurred in 37, 39, and 46 months for cabbage, radish, and
cauliflower in that order (Rodrigo, 1953). In open storage, seeds of
cabbage, cauliflower, and kohlrabi lost viability in 9 months, but seeds
sealed in a desiccator retained their viability for 8 years (Dutt and
Thakurta, 1956). Four lots of swede seeds retained their original
germination for 10 years when stored at 4-percent moisture in sealed
glass bottles (Madsen, 1957). Cauliflower seeds stored similarly re-
tained good viability for 8 years, whereas brussels sprouts seeds sealed
at 4 percent retained their viability well for 16 years. Radish seeds with
4-percent moisture in sealed glass bottles retained their viability well
for more than 8 years. Two lots declined slightly by the end of the 15th
year.
For dry eggplant seeds in a sealed container in a laboratory, loss of
viability occurred in 53 months (Rodrigo, 1953). Tomato seeds with
5.9-percent moisture did not change significantly in viability and vigor
during 36 months of sealed storage at 32° C. With increased seed
moisture, temperature became more critical and tomato seeds lost
viability rapidly above 21° (Asgrow Seed Co., 1954). Potato seeds
stored in sealed containers at temperatures from 0° to that of a
basement room near Greeley, Colo., retained good germination for up
to 13 years (Clark, 1940; Wollenweber, 1942; Stevenson and Edmund-
son, 1950). Germination of seeds at room temperature dropped to 26
percent after 18 years and 17 percent after N years (Wollenweber,
1942).
Use of sealed containers can extend the storage life of lettuce seeds
(Coleman and Peel, 1952; Bass et al., 1962). Air-dry seeds in sealed
containers in a laboratory lost viability in 27 months (Rodrigo, 1953),
but 4-percent moisture seeds retained good viability for 8 years at 21° C
or colder (table 24). Seeds with 7-percent moisture lost all viability in
less than 1 year at 32° and germinated only 4 percent after 4 years at 21°
and 1 percent after 8 years at 10°. Seeds sealed with 10-percent
moisture retained good germination for 8 years when stored at —12 and
—1°. Seeds at 10° kept well for 3 years but deteriorated rapidly
thereafter.
For dry asparagus seeds stored in sealed containers in a laboratory,
loss of viability occurred at 50 months (Rodrigo, 1953). Beet seeds
placed in sealed bottles at 4-percent moisture and stored at room
temperature did not lose viability in 10 years (Madsen, 1957), and

storage in tin cans was superior to storage in waxed cellophane,
chlorinated rubber, or paper bags in linen ones (Coleman and Peel,
1952). Pepper seeds canned with 4-percent moisture or lass stored well
for 36 months at 32° C or lower. As seed moisture content was
increased, the maximum safe storage temperature decreased (Asgrow
Seed Co., 1954). Dry pepper seeds in sealed containers in a laboratory
lost viability at 65 months (Rodrigo, 1953). Okra seeds with 8- to
25-percent moisture content were stored for 10 years in sealed contain-
ers at room temperature of 21°-38° and at 2°-5° in cold storage. The
upper moisture limit for 10 years in cold storage was 12 percent, and a
moisture content as low as 8 percent was required for 7 years of safe
storage at room temperature (Martin et al., 1960).
Safe Moisture Levels for Sealed Storage

It is apparent from this discussion that simply sealing seeds in
airproof and moistureproof containers is not necessarily a satisfactory
packaging procedure for safe long-term storage. The research reviewed
points up the need for careful drying to a moisture level that is safe for
the highest temperature under which the seeds may be stored.
The results of Oregon experiments indicated that the following seed
moisture level percentages can be considered safe for 3 years of sealed
moistureproof storage under moderate temperatures for the following
crops: Alfalfa 6, trefoil 7, clover, perennial ryegrass, soybean, and
sweet corn 8, bentgrass, bluegrass, fescue, timothy, and vetch 9, and
barley, bromegrass, common ryegrass, field corn, oats, rye, and wheat
10 (Ching, 1959).
Moisture content percentages of the following are considered safe for
up to 3 years of sealed storage: Cabbage, cauliflower, pepper, and
tomato 5, celery and lettuce 5.5, cantaloup, cucumber, eggplant, onion,
and watermelon 6, parsley 6.5, carrot and pea 7, beet 7.5, and bean,
lawn grasses, spinach, and sweet corn 8 (Bass et al., 1961).
Koopman (1963) listed the following percentages of moisture as safe
for moistureproof packaging of flower seeds: Ageratum 6.7, Alyssum
6.3, Antirrhinum 5.9, Aster 6.5, Bettis 7.0, Campanula 6.3, Lupinus
8.0, Myosotis 7.1, Nemesia 5.7, Penstemon 6.5, Petunia 6.2, and Phlox
7.8.
The Federal Seed Act and all State seed laws require that all seed lots
offered for sale be retested for germination at specified intervals.
California took the lead in extending the germination test interval for
adequately dried seeds marketed in hermetically sealed containers
(Calif. Dept. Agr., 1973). Federal regulations soon followed California's
lead. The rules and regulations under the Federal Seed Act (U.S. Dept.
Agr., 1968, pp. 17-83) currently provide that the seeds in hermetically
sealed containers shall not exceed the following percentages of moisture

on a wet weight basis:
Moisture-Resistant Materials
Polyethylene Films.—The most extensively used thermoplastic film
is made from an entire family of aliphatic hydrocarbon resins. Com-
mercially available polyethylene resins fall into three groups: (1) Con-
ventional low density types with specific gravity of 0.914 to 0.925 gm
per cubic centimeter, (2) medium density types with specific gravity of
0.93 to 0.94, and (3) high density types with specific gravity of 0.95 to
0.96. Density differences are due to differences in molecular structure.
Molecular structure determines the physical structure of the resins.
Resin properties and extrusion variables determine film properties,
which in turn determine the utility of the film. Physical properties
determine the usefulness of a given film. They include tensile, tearing,
and bursting strength, moisture vapor, carbon dioxide, and oxygen
transmission rates, sealability, elongation, and folding endurance.
Conventional low density films have always been considered more
AGRICULTURE HANDBOOK 506, U.S. DEPT. OF AGRICULTURE;
satisfactory for seed packages than medium and high density films
because of differences it bursting and tearing strength and film elon-
gation or stretch. However, a special medium density film shows
considerable promise. Resin manufacturers and film processors are
continually working to improve bursting and tearing strength and
elongation of medium and high density films.
Medium and high density films tend to show progressively less
permeability to moisture vapor and gases than conventional low density
films. Medium density films show one-half to one-third and high density
films one-fourth to one-fifth the permeability of low density films. A
1-mil low density film tested at 37.8° C and 100-percent relative humid-
ity will permit passage of 1.4 gm of moisture vapor through 100 square
inches of film during 24 hours, a medium density film will transmit 0.7
gm, and a high density film 0.3 gm under the same conditions. A 10-mil
low density film at 37.8° and 100-percent relative humidity will transmit
0.13 gm of moisture vapor per 100 square inches per 24 hours, approxi-
mately one-tenth the amount transmitted by the 1-mil film.
A medium density (specific gravity 0.938) polyethylene film has been
developed that surpasses the performance of conventional polyethylene.
A 7-mil film of this special material has a moisture vapor transmission
rate of 0.10 gm per 24 hours per 100 square inches, which is less than
that of 10-mil conventional polyethylene. This special medium density
film has better tensile properties and greater elongation than conven-
tional polyethylene. Because of its high percentage of stretch, this
medium density film has very good puncture resistance.
Both clear conventional polyethylene and the special translucent,
white, medium density polyethylene films are subject to slow de-
terioration on direct exposure to strong sunlight or ultraviolet radia-
tion. However, deterioration can be retarded by incorporating into the
film carbon black or other pigments, which will absorb the ultraviolet
rays. The special medium density film has a very high resistance to
stress cracking, which has been reported in a few instances with
conventional polyethylene.
Rats and mice sometimes present a problem with conventional pol-
yethylene, but no rodent attack on bags made of the special medium
density material has been reported. Perhaps this material is a solution
to rodent problems.
With a tight closure, such as is produced with a heat seal, both the
10-mil conventional polyethylene and 7-mil medium density polyethyl-
ene bags are almost completely insectproof. Thin films may be pene-
trated by some insects.
Polyethylene films can be laminated to themselves, to other films,
paper, textile fabrics, and fiberboard. Moisture barrier and other
physical properties are improved by laminations. The various proper-
ties of each film included in a laminate are more or less additive. Some
laminated films are completely impervious to various gases and practi-
cally impervious to moisture vapor. Some laminated materials handle
well on automatic packaging machinery and others handle best by hand
depending on the nature of the materials used in the lamination.
Polyester Films.—These films are heat sealable, transparent, flexible
plastic materials, with low moisture vapor, carbon dioxide, and oxygen
transmission rates and great tensile strength. They will not dry out or
become brittle with age because they contain no plasticizer. Polyester
film can be laminated to itself and practically any other material, and its
flexible laminates can be used with most flexible packaging equipment.
A new construction that utilizes a base of light cotton fabric and
metalized polyester film offers easier fabrication, stronger seals, resist-
ance to flex damage, rough handling, and pinholes.
Polyvinyl Films.—These films are heat sealable, deteriorate slowly
in sunlight, and have outstanding tensile strength and tear resistance,
but they provide only moderate protection unless they are laminated to
an effective moisture-barrier material. They heat-seal over a wide
range of temperatures, are ideal for automatic packaging machinery,
and laminate well to paper, foil, or other films. Polyvinyl's resistance to
sunlight and aging indicates that packages will not dry out or become
brittle.
Cellophane.—This family of films is made of regenerated cellulose
and is produced in mere than 100 varieties, each designed for a specific
purpose. Moistureproof types, which have very low moisture vapor
transmission rates, are used for small seed packages. Cellophane alone
does not make a very satisfactory seed package because it becomes
brittle with age or under arid conditions and it breaks easily. Several
firms dealing in flexible packaging materials, however, do produce
combinations of cellophane and polyethylene , which do not become
brittle like cellophane alone and offer fairly good moisture protection.
Polyethylene-cellophane laminates are rather extensively used in seed
packaging as they heat-seal easily and perform well on automatic
packaging machines.
Pliofilm.—Pliofilm is a thermoplastic, rubber, hydrochloride, plastic
film, resistant to ripping, tearing, and splitting. It seals well at low
temperatures, has good moisture-barrier properties, and can be lami-
nated to itself, paper, foils, or other films. Pliofilm can be used on any
package machine designed for flexible film packaging.
Aluminum Foil.—Annealed aluminum foil has a tensile strength of
8.5 pounds per inch of width per mil thickness. It increases in strength
as the gage, or thickness, is increased and as the temperature is
lowered. Tensile strength and resistance to tearing and bursting are
greater for strain hardened foil than for annealed foil of the same
thickness.
Aluminum foil has a low moisture vapor transmission rate, even for
less than 1.5-mil thicknesses, which have tiny perforations called pin-
holes. These seem to be an inevitable result of rolling metal to very thin
gages. Microscopic measurements of all the pinholes in 100 square
inches of 0.4-mil foil gave an estimated area of 0.00004 square inch. A
single hole of this area would transmit about 0.19 gm of water vapor per
24 hours at 37.8° C and 100-percent relative humidity. The number and
size of pinholes decrease with increasing foil thickness. Moisture vapor
transmission decreases also with increasing foil thickness. A 0.35-mil
foil will transmit approximately 0.29 gm of water vapor per 100 square
inches of foil per 24 hours at 37.8° and 100-percent relative humidity, a
0.5-mil foil will transmit 0.12 gm of water vapor under the same
conditions, and thicker films transmit almost no water vapor at all.
Aluminum foil is commonly used in laminations and separately as a
coating and overwrap material for cardboard boxes. Aluminum foil
alone does not make good seed packages, but it can be bonded to other
materials to produce combinations having almost any desired charac-
teristics. Even though thin gages of aluminum foil have some pinholes,
combinations with various supporting materials, such as paper or plastic
films, offer effective barriers to moisture vapor and gas transfer. With
the proper selection of materials, combinations can be produced that
will completely restrict moisture vapor transfer.
Laminations.—Laminations of aluminum foil with other materials
have been satisfactorily used for all sizes of seed packages. Examples of
such lamination., include (1) aluminum foil/glassine paper/aluminum
foil/heat-sealing lacquer, (2) aluminum foil/tissue paper/polyethylene,
and (3) 'craft paper/polyethylene/aluminum foil/polyethylene.
Burlap and cotton can be laminated to paper with asphalt or a
compound latex adhesive to produce a material that will provide good
protection from liquid water but only limited protection from water
vapor. Other constructions utilize such barrier materials as vegetable
parchment, pliofilm, polyethylene, rubber coatings, and foil. Moisture-
resistant multiwall bags usually have a barrier material, such as asphalt
or polyethylene, between the two outer layers of paper. A good asphalt
laminate at 26.7° C and 75-percent relative humidity has a moisture
vapor transmission rate of 0.17 gm per 100 square inches per 24 hours.
Some seed bags are constructed of paper/polyethylene/aluminum foil
laminates. These combinations afford better moisture protection than
either foil or polyethylene used along with paper. Films of cellophane,
pliofilm, polyester, polyvinyl, aluminum foil, and polyethylene are used
alone or in various combinations for seed packages.
PRINCIPLES AND PRACTICES CF SEED STORAGE
Testing Moisture Vapor Transmission of Flexible

Materials
There are several methods of measuring the rate of moisture vapor
transmission through a material. Two methods used in Norway are the
dish method and the method according to Bange (Fornerod, 1963). In
the dish method a strong desiccant is sealed in a special dish by a piece of
the material to be tested. The dish is placed at a controlled temperature
and relative humidity. At periodic intervals the dish is weighed and the
weight change is used to calculate the rate of passage of moisture vapor
through the test material.
The method according to Bange is similar but more accurate for very
slow moisture vapor transmission. The Bange method uses a salt
solution rather than a desiccant in the dish, which is placed in a
measuring apparatus. A strong desiccant is placed in the left pan of the
balance and its weight gain is measured from time to time. From these
measurements the moisture vapor transmission is calculated.
An easy method for an individual to use is to fill a container of the
material to be tested with water, seal it, and place it in a desiccator over
calcium chloride. Moisture penetration is measured by the weight loss
of the package. Conversely, the package can be sealed full of calcium
chloride, then placed in a desiccator over water. In this case weight gain
is used to measure the rate of moisture vapor penetration. No elaborate
special equipment is required for this method of testing moisture vapor
transmission, only a reliable balance for weighing.
From the test data available, not all manufacturers appear to use the
same temperature and relative humidity when testing moisture vapor
transmission of their materials. Some give grams of water per 100
square inches of test material per 24 hours at 37.8° C, with 100-percent
relative humidity on one side and 0 percent on the other side of the
material being tested. Others use the U.S. Bureau of Standards
method, which measures water vapor penetration as grams of water per
100 square inches of test material per 24 hours at 37.8°, with 90-percent
relative humidity on one side and 0 percent on the other. For law
enforcement purposes the Federal Seed Act specifies use of the Bureau
of Standards method.
In the National Seed Storage Laboratory at Fort Collins the moisture
protection of test materials is measured by the change in seed moisture
content over time. Seeds in packages made of good moisture-barrier
materials show little change in seed moisture, whereas seeds in poor
moisture-barrier materials gain or lose moisture rapidly. A simple
method, which does not destroy the seeds, as when making a seed
moisture test, is to weigh the test package when filled and at intervals
thereafter. However, the outside of the package must be free of liquid
water. Also, paper on the outside can hold extra water and give
GRICULTURE HANDBOOK 506, U.S. DEPT. OF AGRICULTURE
erroneous results. The moisture vapor transmission rate can be cal-

culated using the amount of weight change, the time between weigh-
ings, and the surface area of the package. The moisture vapor trans-
mission rate measured by either of the last two methods will be less
than the rate for the same material measured by the U.S. Bureau of
Standards method unless the test package is subjected to the specified
temperature and relative humidity.
Moisture-Barrier Storage
The development of polyethylene and other flexible moisture-barrier
packaging materials prompted studies to determine the value of these
materials as protective packages for seeds. Thin gages of polyethylene,
polyester, and similar materials do not provide very much moisture
protection (Barton, 1949, 1953; Isely and Bass, 1960; Miyagi, 1966).
Storage life was increased by storage in containers made of 5- or 10-mil
polyethylene for the following seeds: Alta tall fescue, Chewings fescue,
corn, creeping red fescue, cucumber, hemp, Highland bentgrass, kenaf,
Kentucky bluegrass, onion, peanut, ryegrass, sudangrass, and wheat
(Anonymous, 1959; Cooper, 1959; Bass, 1960, 1968; Ching et al., 1960;
Toole et al., 1960, 1961; Milligan and Hayes, 1962; and Grabe and Isely,
1969).
Kraft multiwall bags with laminated paper/foil/polyethylene liners
were as effective as tin cans in maintaining a moisture barrier during
storage and shipment of cabbage, soybean, and wheat seeds (Caldwell,
1962). The water vapor transmission rates of polyethylene materials
are inversely proportional to their protective value for seed storage
under tropical, temperate, and warehouse conditions (Ching and Abu-
Shakra, 1965). Packaging materials conta ining aluminum foil provide
good moisture protection (Harrington, 1960u, 1960b, 1963; Lowig, 1963;
Bass and Clark, 1974; and Clark and Bass, 1975).
The need for moisture protection is determined by the storage
conditions and so is the protective value of a barrier material to a
certain extent. A good barrier material, such as a heavy foil laminate,
will provide adequate moisture protection regardless of the storage
conditions; a mediocre material will not give satisfactory moisture
protection when relative humidity is high (table 25).
To provide a rigorous test, we stored crimson clover seeds in a
variety of materials (table 26) in a 20° C night (15 hours) - 30° day (9
hours), water-curtain germination chamber, in which the relative
humidity was 95 to 100 percent constantly. The data (table 25) show
which materials provided satisfactory moisture and germination KT
tection. Seeds in the same materials stored at 10° and 50-percent
relative humidity showed a different moisture and germination picture.
TABLE 25 . —Moisture content and germination of crimson clover seeds packaged in various
flexible materials and stored for 2 years in walk-in germinator held at 20°-30° C and 95- to
100-percent relative humidity (RH) or at 10° and 50-percent relative humidity
TABLE 25. —Moisture content and germination of crimson clover seeds packaged in various
flexible materials and stored for 2 years in walk-in germinator held at 20°-30° C and 95- to
100-percent relative humidity (Rif) or at 10° and 50-percent relative humidity--Con.
I See table 26 for identification of materials.

Seed dead by 6-mo test.
Results of our studies suggest that a foil laminate such as material 10,
11, 14, 15, 16, or 18 (tables 25 and 26), properly sealed, provides as much
moisture protection as a sealed metal can. The data (table 25) also show
that for long-term storage the container must provide adequate mois-
ture protection for the most humid condition to which the seeds could be
subjected, not just the intended storage relative humidity. When
preserving germ plasm, care must be taken to guard against any
eventuality.
TABLE 26 .—Construction and source of 18 flexible packaging materials

tested for suitability as moisture-barrier seed packaged
TABLE 26.—Construction and sources of 18 flexible packaging materi-

als tested for suitablility as moisture-barrier seed packagesl —Con.
Use of Desiccants in Sealed Containers

Although the use of desiccants has received some attention from seed
researchers, it has not received much attention from the seed industry.
The longevity of rice seeds in sealed containers has been increased by
including in the container a drying agent, such as calcium oxide
(quicklime) or calcium chloride (Nakajima, 1927; Kondo and Kasahara,
1941; Ito and Hayashi, 1960). Rice seeds in a closed container with
calcium chloride retained good viability for 9 and 10 years (Nakajima,
1927; Kondo and Kasahara, 1940). Germination of rice seeds having
10-percent moisture when sealed was maintained for 1 year at 40° C
when calcium chloride was enclosed with the seeds. Seed with 16-per-
cent initial moisture was preserved at room temperature with calcium
chloride, and without it all viability was lost (Kondo and Okamura,
1931).
Tobacco seeds sealed with calcium chloride germinated 88 percent
after 11 years and 81 percent after 25 years (Kincaid, 1943, 1958), and
seeds sealed with calcium oxide germinated 87 to 92 percent after
nearly 21 years (Schloesing and Leroux, 1943). Storage with silica gel
improved longevity of timothy seeds (Roberts, 1962), quicklime pre-
served Viola tricolor seeds (Lowig, 1953), but seeds of Zoysia japonica
sealed with calcium chloride lost germination rapidly after 2% years
along with seeds sealed without a desiccant (Radko, 1955). Tempera-
ture had little effect on the longevity of bean, eggplant, lettuce, and
tomato seeds stored over calcium chloride (San Pedro, 1936). Germina-
tion percentages of several crops after 10 years over calcium chloride

were Avena sativa 89, Capsicum annuum 70, Cucumis sativus 80,
Cucurbita pepo 55, Oryza sativa 62, Raphanus sativus 81-89, Solanum
melongena 37, and Zea mays 79 (Kondo and Kasahara, 1940).
Most desiccants do not injure seeds; however, quicklime did have an
injurious effect on seeds of several species, such as Oryza sativa,
Phaseolus radiata var. pendulus (= Vigna radiata), and Vicia sativa
(Nakajima, 1927). Calcium oxide did not improve the longevity of
'Golden Cross Bantam' sweet corn (Edmond, 1959).
Although most of the studies showed that seeds sealed with desic-
cants lived somewhat longer than those sealed without them, their use
has not received wide application in the seed industry because of several
factors. Probably the most important are the added cost of the desiccant
and the larger container required for it. Also, when a package is
opened, the desiccant is soon useless. Because most seeds are stared
commercially only from harvest to the next planting season, the added
expense is not easily justified except possibly in very humid areas
where seeds deteriorate rapidly under normal storage conditions.
Use of desiccants may offer some economic advantage for long-term
storage. However, without the results of a comprehensive study, one
cannot be certain of their value in preserving seed viability and vigor
during long-term storage in sealed moistureproof containers, because
for adequately dried seeds a desiccant may be of no benefit. Such a
study should include several kinds of seeds with a wide range of
moisture levels, stored with different amounts of each kind of desiccant,
and at temperatures from below freezing to 32° C or higher. Desiccants
could be very beneficial when used in conjunction with some of the
flexible packaging materials that have limited moisture-barrier quali-
ties.
MONITORING SEED STORAGE ENVIRONMENT

AND SEED CONDITION
Germination and Viability
Ordinarily seeds are tested for germination and viability by labora-
tories equipped and staffed to determine the quality of seeds. Inexpe-
rienced individuals planning to conduct germination and viability re-
search should receive some training in a reputable laboratory. In the
germination test a seed to be considered as having germinated must
produce a seedling with normal or approximately normal features.
Some kinds of plants produce hard seeds that are considered to be
viable although they do not germinate when tested according to offi-
cially accepted procedures. Sometimes dormant seeds require special
germination test procedures. A viability test aims to quickly detect all

live seeds whether dormant or not. The tetrazolium and the embryo
excision tests are used for this purpose.
The procedures for testing seed germination and viability are found
in the "Rules for Testing Seeds" by the Association of Official Seed
Analysts (1970), special handbooks published by the Association of
Official Seed Analysts, the "International Rules for Seed Testing"
published by the International Seed Testing Association (1966), the
"Rules and Regulations of the Secretary of Agriculture," U.S. Depart-
ment of Agriculture, published in the Federal Seed Act of August 9,
1939, with revisions, and in certain supplementary publications (Grabe,
1970; Justice, 1972). Information concerning the availability of these
rules for seed testing and the special handbooks can be obtained from
the Seed Branch, Grain Division, Agricultural Marketing Service, U.S.
Department of Agriculture, Washington, D.C. 20250.
Seed Vigor
Vigor as applied to seeds is an indefinite term. No definition for seed
vigor has been accepted generally. The methods used to test seed vigor
are just as vague. Neither the Association of Official Seed Analysts nor
the International Seed Testing Association has included methods of
testing seeds for vigor in their procedures. The best known and com-
monly used vigor test is the so-called cold test developed for corn and
subsequently adapted for a few other crop species. One problem with
this test has been the difficulty of standardizing the fungi and soil used
in making the test. Additional information on this test can be obtained
from the Iowa State University Seed Laboratory, Ames 50010, where
the test was developed.
Other procedures used in research include the respiration test, which
measures oxygen consumption and carbon dioxide release, the GADA
(glutamic acid decarboxylase activity) test, various types of stress
tests, rate of seedling growth test, and the tetrazolium test. Each of
these tests has been useful for specific kinds of seeds but has not proved
reliable for a wide range of various kinds of seeds. Woodstock (1973)has
reviewed the literature on vigor tests.
Seed Moisture Content

Practical methods of testing seeds for moisture include oven methods
and electric moisture meters. Basically the oven methods operate on the
principle that the seed moisture is driven off by heating. The difference
in weight of the seeds caused by heating represents the moisture
content. The temperature and time of heating have been reported for
many kinds of seeds. The heating regimes have been developed to

minimize weight changes not produced by moisture.
Large seeds are frequently ground to decrease drying time. Many
seeds are heated to 130° C, whereas others should not be heated above
100° and a few at lower temperatures. A common method is to dry seeds
for 24 hours at 100° to 105°. At the higher temperatures, volatile
materials can be driven off and oils and fats oxidized, both resulting in
weight changes. As the drying temperature is decreased, the drying
time must be increased proportionately. Drying schedules have been
published in the "International Rules for Seed Testing" (1966). These
rules were developed for air ovens that operate at normal atmospheric
pressure. Vacuum ovens with the advantage of reducing drying time
are available. All oven methods have the disadvantage of requiring
much equipment, weighing of materials, and considerable time for
testing. However, they are the more accurate of the practical methods.
Electric moisture meters have an advantage in speed over all other
practical methods. Most electric moisture meters are based on meas-
urements of either conductivity or the dielectric properties of the seed.
Although these measurements vary with the moisture content of the
seeds, they are affected to some extent by other factors. Calibration
charts have been developed for many kinds of seeds. Calibrations
should be based on the testing of relatively large numbers of samples,
covering a wide range of moisture contents, sources of samples, and
years of harvest.
With most moisture meters, the test can be completed within 1 or 2
minutes. Disadvantages are relatively high cost of equipment, need for
careful, painstaking calibration for each kind of seed, and in some cases,
especially for very wet and very dry seeds, failure of the equipment to
be highly accurate. For more information on methods of testing seeds
for moisture content, see Hlynka and Robinson (in Anderson and
Adcock, 1954), Hart et al. (1959), and Zeleny (1961).
Fungi and Bacteria

Methods of testing seeds for the presence of seedborne micro-orga-
nisms are given in the "International Rules for Seed Testing" (1966).
For additional information, see Muskett (1948), de Tempe (1961),
Naumova (1970), Baker (1972), and Neergaard (1973).
Relative Humidity
A psychrometer is commonly used to measure relative humidity.
Sling psychrometers are inexpensive, reasonably accurate, and easy to
use. This instrument consists of a dry-bulb thermometer, which
measures the ambient temperature, and a wet-bulb thermometer,
which measures the reduced temperature caused by the cooling effect of

the evaporating liquid. This cooling effect is in proportion to the relative
saturation of the air being tested. Recording psychrometers have the
advantage of providing a record at any time over the interval covered
by the clock and chart. Both temperature and relative humidity are
recorded by the hygrothermograph. This instrument requires special
attention when changing the charts. The directions for its use should be
followed very closely. Recording units are available that use' electrical
resistors to measure humidity and thermistors to measure temperature.
Although expensive, they are very accurate. Major changes in relative
humidity can be detected with cobalt chloride cards. At different
positions on the card, the color changes as the relative humidity
progressively increases.
Temperature
Thermometers, like psychrometers, are made in several forms. When
a high degree of accuracy is not required, simple, inexpensive stem-
type thermometers can be used to measure the temperature of the air
or the mass of seeds. Although some or mos of the thermometers of
this type may be accurate, they must be removed from the seed mass
before taking the reading. By the time the reading is taken, the
mercury column may have changed, and thus it gives an erroneous
reading. Recording thermometers, with or without a sensing clement,
are available. Of course, these instruments provide a continuing record,
which can be cited for future reference if desired. Multipoint recorders
provide essentially the same information, but each instrument can be
equipped with several sensing units, or thermocouples, allowing the
monitoring of several locations simultaneously. Thus, hot spots in a
mass of seed or grain may be detected with a thermocouple that
conceivably could go undetected if the other temperature sensing and
recording instruments were used. Thermographs, recording thermom-
eters, and multipoint recording units are rather expensive. Their use
may or may not be justified depending on the degree of accuracy
desired, chance of product heating in storage, extent of business
operation, and frequency of use.
Storability of Seed Lots

Considerable research, mostly at the Mississippi State University
Seed Technology Laboratory, has been conducted to develop a practical
me hod for predicting relative storability of seed lots, especially those
to stored for future use. Although several potential tests were
devised and evaluated in this research, two were more successful than
others—the accelerated aging test and the modified aging test.
The following information has been taken from reports by Delouche

et al. (1967, 1968), Delouche and Baskin (1971), and Goff (1972). "In the
accelerated aging test, the germinative responses of representative
samples of seed lots are determined after exposure to conditions of
temperatures of 40° to 45° C and 100-percent relative humidity for
varying periods of time up to -204 hours. The modified aging test
measures the germinative responses of samples after they have been
held at 30° and 75-percent relative humidity for various periods of
time."
Optimum accelerated aging conditions and treatment times in the 2
tests are given for 16 kinds of seeds in table 27.
TABLE 27. —Optimum accelerated aging conditions and treatment

times in 2 tests for predicting seed storability of 16 crops'
We do not claim that these tests will accurately predict the storability
of all seed lots of a given crop species. The principles involved do offer
the seedsman a means of detecting some seed lots that should not be
stored for extended periods of time. More accurate predictions could be
made if the seed lots were stored under controlled temperature and
relative humidity rather than under ambient conditions.
AGRICULTURE HANDBOOK 5u6, U.S. DEPT. OF AGRICULTURE
The researchers who developed the accelerated aging test of pre-

dicting seed storability are confident that the method or modifications
will ultimately be perfected so that storability of seed lots can be
predicted with considerable accuracy. Additional information on these
test procedures can be obtained from the Seed Technology Laboratory,
Mississippi State University, State College 39762.
Detection and Identification of Fungi and Insects

Ordinarily the manager or superintendent of the storage warehouse
must monitor his seed stocks. It is his responsibility to check for
evidence of fungi and insects. Their presence may be detected either by
observing their growth on or in the seed In ass or by development of hot
spots. If hot spots are detected, the cause should be ascertained
immediately. The cause may be traced to molds, insects, or both.
Specialists may be required to identify molds or insects. The State
agricultural colleges and universities and State departments of agricul-
ture frequently will make simple identifications of occasional specimens
for residents. Depending on their background, some county agricultural
agents may perform this service. Seed companies often employ a plant
pathologist. Some official and commercial seed testing laboratories
routinely conduct seed health tests.
SOME PRACTICAL INFORMATION FOR STORING

AND TRANSPORTING SEEDS AT AMBIENT
CONDITIONS
Examples of Storability of Different Plant Species
These much variability in the lifespan or storage life of seeds of
different plant species. The lifespan of seeds of certain aquatic and
woody plants is very short, whereas seeds of some leguminous species
and a few other kinds retain viability for over a hundred years. Only the
practical storage life of cultivated species of field, vegetable, herb, and
Cower seeds is discussed here, not species with extremely short or
extremely long storage lives. Seed longevity per se is not included. The
information here relates to the retention of viability of most individual
seeds in a lot, not the elapsed time before the last viable seed in a lot or
sample is dead.
Species for which information is available have been listed. The data
can be used as guidelines for closely related species not given. The
information is rather scanty, some of it is outdated, and most of it has
been obtained under nonuniform storage and test conditions. This
obviates accurate comparisons.
As indicated by the column heading, "Relative Storability Index," in

the tabulation, the data do not refer to specific years of storage. In
general, the authors classified in category 1 those plant species of which
50 percent or more of the seeds can be expected to germinate after 1 to
2 years of storage under favorable ambient conditions at latitudes of
approximately 35° to 48° N., in category 2 after 3 to 5 years, and in
category 3 after 5 or more years.
Examples of relative storability of field, vegetable, herb, and flower
seeds of high viability and vigor are as follows:
The following references were used in compiling the list: Bodger

Seeds Ltd. (1932), Filter (1932-33), Goss (1937), Harrington (1972),
James et al. (1964), Madsen (1962), Owen (1956), Pritchard (1933),
Sifton (1920), Ullmann (1949), Welton (1921), and Wheeler and Hill
(1957).
Climatological Data Pertinent to Seed Storage

Although the relative humidity and temperature of the storage
environment can be controlled artificially, the cost of providing such
control in large storage areas is usually not economically feasible. Thus,
most seeds stored in North America are kept at ambient temperatures
and relative humidities. Storage as used here includes holding of seeds
for processing, shipment, or sale, as well as the period of shipping.
Because of the wide range of climatic conditions in the United States,
its territories, Puerto Rico, Canada, and Mexico, climatological data
useful to seedsmen, seed merchants, and farmers could be provided in
maps; however, weather maps give less specific information than tabu-
lar data and may be difficult to interpret.
The data in table 28 are relatively longtime averages and should
provide reliable guidelines for persons planning to store seeds at
ambient conditions at the indicated locations. Data are given for Jan-
uary, April, July, and October, the first month of each season, as well as
annual averages. Locations have been selected to give a fairly repre-
sentative cross section of the area covered. For locations not shown,
data from the nearest station should be used or adapted, based on
known facts about the local climate. For example, if one is going to store
seeds at ambient conditions in Dayton, Ohio, from July to January, he
would probably check the conditions at Cincinnati and Columbus, Ohio,
as well as at Indianapolis, Ind. Six-month averages for these three
locations for July and October are temperature 19° C and relative
humidity 70.1 percent. For practical purposes, these values would be
close enough for Dayton. If the seed is to be stored for a year, the
annual averages for the three locations should be used, as 12° and
71.7-percent relative humidity. Other adaptations may be necessary for
other locations, especially where relatively great climatological dif-
ferences occur within short distances, as in mountainous areas or near
large bodies of water. Annual averages should be used with great care
because of extreme seasonal differences that can be masked in them.
These data provide a guide for drying carryover seed and storing it in
moisture-barrier containers. It should be remembered that basement
rooms, unless well ventilated, always have a higher relative humidity
than aboveground rooms and can be potentially dangerous as seed
storage areas.
Another possibility is to obtain original and up-to-date information
from the National Weather Records Center, U.S. Department of Com-
merce, NOAA, Asheville, N.C. 28801. This Center can provide current,
more complete, and longtime averages of temperatures, relative hu-
midities, and several other weather and climatological data for many
stations not shown in table 28.
Relation of Storage Conditions to Intended Storage

Periods
The first question asked of seed storage specialists is "How should I
store my seed?" For an adequate answer the specialist must know
where and what kinds are to be stored for how long. In arid areas many
kinds of seeds can be stored for 5 years or longer under ambient
conditions provided the temperature is not too high. In temperate
regions the same kinds of seeds may keep only 2 or 3 years, whereas in
the humid subtropical and tropical regions viability may be lost in a few
months or even weeks at ambient conditions.
A useful guide for commercial seed storage for up to 5 years is
Harrington's (1960d) thumb rule that states "the sum of the tempera-
ture in °F and the percent relative humidity should not exceed 100." For
example, various kinds of seed stored at 50° F (10° C) and 50-percent
relative humidity showed no loss in viability during 5 years of storage
(James et al., 1967). Comparable results can be obtained by drying seed
to 5-percent or lower moisture content and storing it in sealed mois-
ture-barrier containers at ambient temperatures up to 32° C. At higher
ambient temperatures some refrigeration is desirable.
TABLE 28.—Climatological data for 183 weather stations in the United States, its territories, Puerto Rico, Canada,
and Mexico1
TABLE 28. —Climatological data for 183 weather stations in the United States, its territories, Puerto Rico, Canada,
and Mexico1—Continued
and Mexico1---Continued
See footnotes at end of table .
TABLE 28.—Climatological data for 183 weather stations in the United States, its territories, Puerto Rico, Canada,
1 Data from (1) "Local Climatological Data, Annual Summary With Comparative Data," 1972, and (2) "Climatic Atlas of the United States," 1968,
both published by National Oceanic and Atmospheric Administration, U.S. Department of Commerce; (3) "Atlas of American Agriculture," U.S.
Department of Commerce, 1928; (4) "U.S. Naval Weather Service World-Wide Airfield Summaries," v. VII (Central America), February 1968; and
(5) Meteorological Branch, Canada Department of Transport.
2 Each average relative humidity, except one, represents the average of 4 readings taken at approximately 12 p.m., 6 a.m., 12 m., and 6 p.m. For
Milford, Utah, the average represents readings taken at 11 a.m. and 5 p.m., the only data available. Each fraction resulting from averaging
temperatures and relative humidities is rounded to the nearest whole number; 0.5 is raised to the next number.
Using the correct seed moisture content for sealed storage is very
critical as seed sealed with too high a moisture content may lose
viability more rapidly than air-dry seed in open storage at the same
temperature. For very long-term storage, seed should be dried to
5-percent or lower moisture content, sealed in moisture-barrier con-
tainers, and held at —17.8° C, whereas storage at 4° and 50-percent
relative humidity in open containers is satisfactory for many kinds of
seed for 10 years or longer.
There are, of course, exceptions, as some kinds of seeds cannot
endure freezing and others cannot tolerate dehydration without losing
viability. The cost of storing a given quantity of seed increases rapidly
as the storage conditions become more exacting. Thus, the storage
facility should be related to storage time, commensurate with the needs
for maintaining satisfactory levels of seed viability and vigor. All
storage structures must be designed to protect the seeds from insect
and rodent infestations.
It should be emphasized that the intended storage period is only one
of several items to be considered when deciding on storage conditions
for a particular lot of seed. The more significant of these are seed
factors, storage environment, effect of pests, and storage structures.
Care of Seeds in Transit

Storage Principles Applicable to In-Transit Seeds
As pointed out by Harrington (1972) and others, seeds are in storage
from the date of their physiological maturity until they are planted.
Thus, seeds in transit are, in fact, in storage. As used here, the
expression "in transit" includes not only the time the seeds are being
moved from one location to another but also the time they are awaiting
shipment, whether in a warehouse, railroad car, truck, airplane, boat,
or on a dock awaiting further consignment. While in transit the seeds
are subject to the same storage principles as seeds in warehouses. The
principal differences between warehouse storage and in-transit storage
are the relatively fast changes in the seed environment that can result
from modern transportation and the failure of personnel to foresee or
predict all the hazardous conditions to which the shipment may be
subjected. The two principal envionmental factors that determine
whether a seed shipment will arrive safely at its destination are the
temperatures to which the seeds are exposed and the seed moisture
content, which is controlled by the relative humidity of the air or
protective packaging.
Historical Background
Moodie ( 1925) reported on the effect of temperature on seeds shipped
from Australia to England and from England to Australia. Since the
temperatures used aboard ship were not precise and moisture content
of the seeds was not controlled, his results are not completely convinc-
ing. From his experiments Moodie concluded that seeds stored under
cool conditions with little variation are likely to retain viability better
than when stored under warm conditions with extensive variations in
temperature. von Degen and Puttemans (1931) shipped predried and
nondried seeds of several species of field crops and vegetables between
Hungary and Brazil. Their results proved conclusively the beneficial
effect on germination of predrying the seeds before shipment and
maintaining low moisture content during transit.
Foy (1934), working at Palmerston North, New Zealand, and Kearns
and Toole (1939), working at Washington, D.C., proved conclusively
through cooperative studies that Chewings fescue seed could be shipped
between New Zealand and the United States and between New Zealand
and England without serious loss of viability by predrying the seed
before shipment and avoiding exposure to high temperatures and
humidities. Their carefully conducted studies, which included seeds
exchanged between the three countries and seed stored at Washington,
Palmerston North, and Cambridge, England, were carried out on only
one kind of seed, Chewings fescue. However, the basic principles
relating to seed moisture content as well as relative humidity and
temperature during shipment govern the maintenance of viability of
transoceanic shipment of various kinds of seeds. The results are so
convincing that very little research of this nature has been reported
since.
The deleterious effects of high temperature, high relative humidity,
or both are mitigated by the length of time the seed will be under a
particular set of conditions. For example, a shipment of seed from Cape
Town, South Africa, direct to New York by modern freighter may
arrive at its destination without any loss in germination. On the other
hand, a similar shipment by tramp freighter, which would spend up to 6
weeks stopping at Belem and Manaus, Brazil, might arrive at New York
with significantly reduced germination. Both cargoes in this hypotheti-
cal example passed through tropical waters; however, the extra time
the seed was exposed to hot humid conditions in the Tropics could be
disastrous for the seed.
Some Hazards To Be Avoided
A person planning to ship seeds should anticipate the various hazards
to seeds during transit and while in storage immediately before and
after shipment. These hazards will vary depending on the method of
transportation—whether by truck, railroad car, ship, or airplane. Rapid
changes of temperature or relative humidity resulting from the move-
ment of seeds between zones of different temperatures and humidities
can create hidden problems. Seeds chilled during air transportation are
likely to be covered with condensed water after larding of the plane at

warmer temperatures. Under some conditions this surface moisture can
be absorbed by the seeds rather rapidly. Proper storage immediately
after landing will obviate this problem.
Ching ( 1959) pointed out that under simulated shipping and storage
experiments Highland bentgrass seed absorbed water rapidly under
tropical conditions and lost its viability within a few weeks. It is
noteworthy that Ching found seed stored under arctic conditions tended
to absorb water gradually during prolonged storage wild consequently
gradually lost viability. She recommended that shippers not send high
moisture seed, such as that stored at high humidities, during the winter
into areas where it would be exposed to high temperatures.
Harrington (1972) pointed out the peril of seeds becoming overheated
when shipped in boxcars placed on a railroad siding in a warm or hot
climate for several days. The boxcar acts as a heat trap whereby very
high temperatures may be reached. This hazard is greatest with high
moisture seeds.
Shipping seeds with borderline to high moisture contents is no doubt
the greatest of all hazards. Christensen ( 1969) stated that much grain is
shipped and stored at moisture contents slightly above safe levels at
anticipated temperatures. When the temperature rises purposely, ac-
cidently, or unexpectedly, respiration increases, storage molds appear,
and spoilage results. This is more likely to occur in grain shipped in bulk
than in seeds ordinarily shipped in bags or smaller containers.
Still another hazard is the possibility of shipping seeds as a part of a
mixed cargo. When considering this possibility, the shipper should be
sure that no chemicals that are deleterious to the seeds, such as certain
herbicides, are included in the cargo. Any substance that might increase
the concentration of oxygen should be excluded as oxygen hastens seed
deterioration. There is no objection to inclusion of the inert gases, such
as hydrogen, nitrogen, and carbon dioxide, which tend to retard de-
terioration.
Another hazard is the possibility of the seedsman or shipper failing to
carefully select the seed lots. Seeds of different crop species vary as to
their storability. Oily seeds deteriorate more rapidly than starchy
seeds. Storability differences may exist among cultivars of some crop
species and also among different crops of the same cultivars or species.
Such information when available should be used in choosing seed lots for
shipment. If seeds are to be shipped long distances under unfavorable
or questionable conditions, only fresh seeds of high viability and vigor
should be chosen.
General Recommendations
(1) Carefully select the seed lot for shipment. Consider the general
guidelines given previously.
(2) Carefully consider such factors as the shipping quality of the

seed lot selected, method of shipment, distance to be shipped, time in
transit, and climatic zones through which the seed will pass.
(3) If the seed is not of safe moisture content for shipping, reduce
moisture to a safe level by an acceptable drying method.
(4) If feasible, ship the seed under controlled temperature and
relative humidity to assure viability. If not feasible, reduce the seed
moisture content to a safe level and ship in moisture-resistant or
moistureproof containers (Caldwell and Bunch, 1961). See the sections
on "Seed Storage Structures" and "Packaging and Packaging Materi-
als."
THEORIES REGARDING SEED DETERIORATION

We have pointed out how growing, harvesting, processing, and
storage conditions affect seed longevity. We have also discussed ways to
slow down the rate of seed deterioration, but we have not discussed
what occurs inside a dry, sound, disease-free seed to cause it to weaken
and the. Several theories based on genetic and physiological principles
have been proposed.
Changes in Protein Structure

Ewart (1908) theorized that seed longevity depends not on available
food reserves but on how long the proteid molecules, into which
protoplasm disintegrates when drying, can recombine into active pro-
toplasm with the absorption of water. According to this reasoning,
protein molecules should disintegrate excessively when seeds are dried
to very low moisture levels. Struve,10 however, concluded that cern
dried to less than 4-percent moisture, sealed in nitrogen, and stored at a
low temperature would keep indefinitely. Nutile (1964b) found that
drying vegetable seeds to 0.4 percent caused some damage to carrot,
celery, eggplant, pepper, tomato, and Kentucky bluegrass seeds but not
complete loss of viability. Seeds of cabbage, cucumber, lettuce, onion,
and Highland bentgrass sealed with 0.4-percent moisture were not
injured.
Crocker (1938) suggested that protein coagulation caused loss of
viability. Later he (1948) reported that his protein coagulation theory
was too general because of the many kinds of protein in embryos and
because his studies did not show which protein coagulates with aging.
Depletion of Food Reserves

The theory that depletion of food reserves for the embryo causes
seeds to die did not persist long because it was soon evident that many
dead seeds still contained ample food reserves. Some Zea mays L.
seeds, more than 700 years old, found in the Mesa Verde cliff dwellings,
appeared sound visually, yet not a single viable seed was ever found
among them. Oxley (1948) suggested that exhaustion of an unnamed
organic compound results in loss of seed viability. Harrington (1960b)
reasoned that a seed may have an adequate food supply and still die
because of a breakdown in the food transport system. Seed moisture
content may be high enough for respiration but too low to transport
food from the reserve supply to the embryo (Harrington, 1967). Al-
though food reserves are usually not entirely depleted when seeds die,
various changes may have occurred, such as increased acidity (Zeleny
and Coleman, 1938, 1939; Milner and Geddes, 1946) and decreased
lipids and proteins (Pomeranz, 1966; Ching and Schoolcraft, 1968;
Koostra and Harrington, 1969).
Development of Fat Acidity

The development of fat acidity in seeds has been shown to accompany
death. Germination declined 8 percent and fat acidity increased 14 units
when 8- to 9-percent moisture content soybeans were stored for 700
days. With increased seed moisture content, germination dropped rap-
idly and fat acidity increased sharply (Holman and Carter, 1952). A
drop in the germination of wheat seeds was accompanied by an increase
in fat acidity (Kelly et al., 1942). Increased fat acidity was also a major
cause of corn seed deterioration (Zeleny and Coleman, 1939). Fat
acidity of peanuts increased only after the stored seeds were dead
(Davis, 1961). Although fat acidity values have been established for
grain showing little or no deterioration (Baker et al., 1957), such values
may not constitute a reliable index of viability (James, 1967).
Enzymatic Activity
Attempts have been made to use enzymatic activity as a measure of
seed viability. However, only a few of the many enzymes in seeds have
been investigated. Early work with enzymes dealt principally with
catalase activity. Crocker and Harrington (1918) found catalase activity
in dead Johnsongrass and yet reported a relationship with viability.
They could not establish such a relationship with seeds of Amaranthus
retroflexus. Davis (1926) showed a relationship between catalase ratio
and viability of lettuce seeds, but the range of his study was not wide
enough to establish a linear relationship. Leggatt (1929-30) obtained a
correlation between catalase activity and germination of wheat seeds.
Because of the limited number of species studied and the inconsistent
results, the relationship between catalase activity and seed viability

appears questionable.
Phenolase activity also does not appear to be an indicator of viability
(Davis, 1931). For wheat, Davis found a relationship with germination
but not age, and for oats a relationship with age but not germination.
Throneberry and Smith (1955) reported that malic dehydrogenase ac-
tivity was more closely correlated with germination percentage than
was alcohol dehydrogenase and cytochrome dehydrogenase activity.
They believed inactivation of these enzymes was not a major cause of
viability loss. Linko and Sogn (1960), Bautista and Linko (1962), and
Grabe (1964) demonstrated a close relationship between glutamic acid
decarboxylase activity and viability. James (1968) found that the corre-
lation coefficient between germination percentage and glutamic acid
decarboxylase activity for the average of all bean varieties studied
compared favorably with the coefficients reported by Grabe (1964); for
corn, the coefficients for individual varieties were variable. James
(1968) stated that the inconsistencies of the correlation coefficients
among the cultivars used definitely limit glutamic acid decarboxylase
activity as an indicator of viability of bean seeds. Because of the
conflicting reports on the relationship of enzymatic activity to seed
deterioration, much research remains to be done in this area.
Chromosomal Changes
In 1901 De Vries (in Kostoff, 1935) observed that old Oenothera
lamarckiana seeds produced more abnormal plants than did fresh seed.
More recently chromosomal changes have been reported in old seeds of
a relatively large number of species: Crepis spp. (Navashin, 1933;
Navashin and Shkvarnikov, 1933; Navashin and Gerassimowa, 1935),
corn (Peto, 1933), onion (Nichols, 1941, 1942), sugar beet (Lynes,
1945), barley, pea, rye, and wheat (Gunthardt et al., 1953), Datura spp.
(Avery and Blakeslee, 1936; Blakeslee et al., 1942; Blakeslee, 1954),
and barley, broadbean, and pea (Abdalla and Roberts, 1969a).
Although numerous compounds are known to induce mutations, few
of them are normally found in seeds. Mutagenic compounds found in
plants include adenine, degradation products of adenine, uracil, thy-
mine, adenosine desoxyribonucleic acid, and ribonucleic acid (D'Amato
and Hoffman-Ostenhof, 1956). The degrees of mutagenic action given
by these authors are (1) the lethal zone, where the accumulation of
mutagens becomes toxic and kills the seed, (2) the narcotic zone, which
affects the spindle mechanism, and (3) the subnarcotic zone, in which
the mutations occur. The mutagenic or chromosomal aberration theory
is supported further by the fact that (1) extracts from old seeds induce
mutations in fresh seeds, (2) the mutation rate increases with age, (3)
spontaneous mutations in dormant seeds become evident in the chro-
mosome presplit phase and in adult plants, and (4) reactions of root and
shoot tips in old seeds closely parallel the reactions of the same kind of
seeds when treated with X-ray. Apparently mutagens do not develop
under good storage conditions, as all observations have been on seeds
affected by high temperature, relative humidity, or both.
Blakeslee's (1954) work with Datura demonstrated that age alone was
not responsible for the development of mutations. He found that
mutations developed in seeds at room temperature. The mutation rate
for seeds buried in the ground for 39 years was extremely low. James
(1967) suggested that seeds presently housed in the National Seed
Storage Laboratory at Fort Collins, Colo., may hold the ultimate
answer to the question of the relationship of mutations to seed age.
However, the answer will not be determined soon because the storage
conditions there are such that respiration in the stored seeds appears to
be near the minimum.
Membrane Damage
According to Villiers (1973), the immediate damage rendering aged
seeds incapable of germination is extranuclear. Free radical damage to
membranes and enzyme systems could affect essential metabolic proc-
esses when the seeds become imbibed for germination. Harrington
(1973) stated that seeds dried to 4- to 5-percent moisture content appear
to deteriorate slightly faster than seeds with 5- to 6-percent moisture,
probably from lipid autoxidation damage. Unsaturated lipids in seed
cells may break, producing two free radicals, which can react with other
lipids, destroying the structure of cell membranes. In imbibed cells
tocopherols made by enzymes combine with free radicals rendering
them harmless. Since enzymes are inactive at low seed moisture con-
tents, the free radicals produced nonenzymically become destructive
when the tocopherols that were present When the seeds were dried are
depleted.
Respiration
The theories for seed deterioration, except possibly fat acidity, are
related to respiration. Respiration increases in proportion to the
amount of moisture in seeds, it is very low at moisture contents
between 4 and 11 percent (Ba v, 1940; Harrington, 1963). Respiration
rates up to about 59° C are also directly proportional to temperature.
With high temperatures and high moisture contents, seeds lose viability
very rapidly, usually in less than 3 months at 32° and 90-percent relative
humidity. Seeds stored at temperatures below 10° and at low relative
humidities will remain viable for a long time. According to James (1967) ,
peanut seeds recognized as short lived were held at the Southern

Regional Plant Introduction Station, Experiment, Ga., for 8 years at
10° and 50-percent relative humidity without a significant decrease in
viability.
Summary
What really causes a seed to deteriorate? We examined the theory of
changes in protein structure and found that no specific changes have
been pinpointed. Depletion of food reserves is not a very sound theory
as almost all seeds contain an abundance of food long after the embryo is
dead. Although the development of fat acidity has been shown to
accompany the death of a seed, such development has not been firmly
estalished as the cause of death. No specific enzyme activity change has
been proved to cause the death of seeds. Chromosomal changes have
been noted in various kinds of seeds; however, no one has yet proved
conclusively whether such changes are really the cause of deterioration
or are just a result. Membrane damage, though not specifically es-
tablished as the cause of deterioration, is definitely associated with it.
Although several theories have been proposed, none satisfactorily
explains how seeds deteriorate. Even though the process of deteriora-
tion is not clearly understood, the methods for preventing it are well
established. For more detailed information on theories of seed de-
terioration, see Streeter (1965), Roberts (1972), Abdul-Baki and An-
derson (1972), and Heydecker (1973).
OLD AND ANCIENT SEEDS

Ewart (1908) wrote as follows about the longevity of seeds:
"Probably few sections of human knowledge contain a larger per-
centage of contradictory, incorrect and misleading observations than
prevail in the works dealing with this subject, and, although such fables
as the supposed germination of mummy wheat have long since been
exploded equally erroneous records are still current in botanical physi-
ology. In addition, there are considerable differences of opinion as to the
causes which determine the longevity of seeds in the soil or air. The
works of de Candolle 1832, 1846, de Candolle and Picket 1895, Duvel
1905 and Becquerel 1934 are the most accurate and comprehensive
dealing with the question. The subject is still, however, in an incom-
plete and fragmentary condition."
The principal objectives here are twofold. First, we establish that
very old seeds in some instances have retained vitality over a long
period, even more than a century. We differentiate (1) crop seeds
known to have survived for relatively long storage periods, (2) seeds of
native plants known to have survived longer than crop seeds, and (3)
circumstantial evidence of seed survival for a few to several centuries.
The second principal objective is to analyze the authenticity of so-called

mummified seeds or grains.
Maximum Known Survival of Crop Seeds

Unless crop seeds are kept under favorable storage conditions, they
lose viability within a few years. The information in the section on
"Examples of Storability of Different Plant Species" confirms this
broad-based general statement. On the other hand, there are several
authentic records of cereal crop seeds surviving up to 32 years. Auf-
hammer and Simon (1957) reported survival of barley and oat seeds for
123 years (table 29). Authentic records show that seeds of some of the
leguminous crops have survived up to 81 years (table 29).
All the results in table 29 are regarded as authentic. Some data have
been obtained from planned experiments and others from tests on seeds
stored in museums, cupboards, or laboratories under such conditions
that the history of the seed was known.
Maximum Known Survival of Primarily Weed and

Native Plant Seeds
Mature seeds of many weeds and native plants have dormant em-
bryos, a condition that retards germination. In fact, seeds do not
germinate until they undergo physiological and biochemical changes
that permit afterripening. A dormant or quiescent embryo appears to
enhance longevity. Also, seeds with hard coats, which restrict imbibi-
tion of water and exchange of gases, are among those with the greater
lifespans. Many seeds of weeds and native plants are in this group,
which includes such plant families as the Convolvulaceae, Leguminosae,
Malvaceae, and Nymphaeaceae.
Plant species whose seeds have long lifespans are listed in table 30.
The data were obtained from seeds kept or stored dry with known
histories and from seeds buried in soil by planned experimentation.
Seeds Stored Dry
Perhaps the earliest authentic records of tests on the continued
vitality of air-dry seeds are those of Alphonse de Candolle (1846). He
(1832) tested seeds of different species harvested in 1831 and kept them
all air-dry until May 1846 (15 years), when he planted 20 seeds of each
species. There were 368 species representing 53 families. Five out of 10
species of Malvaceae, 9 out of 45 species of Leguminosae, and 1 out of 30
species of Labiatae showed some germination.
Ewart (1908) published the results of tests of over 1,000 species of
seeds, which he found locked in a cupboard in the botanical laboratory at
Melbourne, Australia. The seeds had been sent from Kew Gardens,
TABLE 29. —Examples of documented long lifespans of some common crop seeds when maintained under favorable
survival conditions
TABLE 29. —Examples of documented long lifespans of some common crop seeds when maintained under favorable
survival conditions—Continued
TABLE 30. —Records of seeds 75 years or older with some germination
England, in 1856 for the University Gardens, but they had been put
away in a dark, dry closet, where they had remained unopened for 50
years or longer. He soaked the seeds and placed them on moist filter
paper in glass dishes in germinators. Seeds that did not swell after 1 or
2 days in water were either filed or treated with concentrated sulfuric
acid. After washing, the seeds swelled readily. Out of 1,400 kinds
tested, Ewart (1908) found 58 that retained their vitality after 50 years
or more in storage. He also tested seeds from another source that
varied in age up to 105 years. Of these seeds, Goodia latifolia and
Hovea linearis germinated 8 and 17 percent, respectively. Of the 58
kinds surviving 50 years or more, 36 were in the Leguminosae, 4 in the
Malvaceae, 2 in the Euphorbiaceae, and the remainder in miscellaneous
plant families. Dry seeds of many leguminous species are frequently
hard. The results of some of Ewart's tests are in table 30.
Becquerel (1934) had access to a batch of old seeds in a storage room
in the National Museum of Paris. Thes seeds were collected from 1819
to 1853. He conducted germination tests on them in 1906 and again in
1934. Since they were all hard-coated seeds, they required special
treatment. They were sterilized, the coats were broken, and they were
put in tubes under sterile conditions at 28 0 C to germinate. The seed
stock was considered so precious that only 10 seeds of each kind were
used for the test. For the 1934 test he obtained from another source
about 20 seeds of Cassia multijuga, which were collected in 1776. Only
two of these seeds were tested. Table 30 shows the results obtained
with six species that germinated after 75 or more years. All these seeds
were Leguminosae. The seeds of C. multijuga germinated after 158
years of storage. Becquerel believed the long lifespan of all these seeds
was made possible by impermeability of the coats, which prevented any
exchange of gases or water between the interior of the seed and the
outside atmosphere, and by the high degree of desiccation (2- to
5-percent moisture) and absence of oxygen in which the embryos
existed within the hard coats.
Turner (1933) tested the viability of old seeds from different sources
kept in loosely corked bottles in a museum. Of nine species listed in his
publication, seven retained viability in excess of 75 years (table 30). All
were Leguminosae.
Schjelderup-Ebbe (1936) tested the viability of 1,254 batches (nearly
as many species) of seeds stored in bottles or paper bags for 34 to 112
years. Seeds of 3 species survived for 50 to 59 years, 10 for 60 to 69
years, 3 for 76 to 74 years, and 2 for over 75 years (table 30). Of 54
entries in the author's table, 19 species germinated after 50 or more
years of storage. One species belonged to the Cannaceae, 13 to the
Leguminosae, 4 to the Malvaceae, and 1 to the Convolvulaceae. Seeds of
some species in all these plant families produce hard seeds.
Buried Seed Experiments

The first planned experiment in America to test the longevity of seeds
buried in soil was begun in Michigan. In the fall of 1879, Beal (1885) of
the Michigan Agricultural College began an experiment to determine
the longevity of seeds of 23 kinds of weeds growing near East Lansing,
Mich. Fifty freshly harvested seeds of each of the 23 kinds (2 woody and
21 herbaceous) were mixed with sand and placed in pint bottles. The 20
bottles were buried. They were inclined at a 45° angle 18 inches below
the soil surface on a sandy knoll of the college campus.
The plan was to dig up one bottle every 5 years and test the seeds for
germination. This schedule was followed until 1920, when it was decided
to remove bottles at 10-year intervals to extend the experiment. Seeds
of Brassica nigra and Polygonum hydropiper showed some germination
for the last time in the 50th year (Darlington, 1931). Seeds of Oenothera
biennis, Rumex crispus, and Verbascum blattaria germinated in each
succeeding test until the 80th year (Darlington and Steinbauer, 1960).
As indicated in table 30 for Beal's experiment, V. btattaria was the only
species that germinated in the 90th year (Kivilaan and Bandurski,
1973).
Duvel (1905) set up a more elaborate buried seed experiment in 1902
at Arlington, Va. It was designed to test the longevity of 107 species of
crop and weed seeds. The seeds were mixed with sterilized soil, placed
in porous flowerpots, covered with inverted flowerpot saucers, and
buried at depths of 8, 22, and 42 inches. After 20 years' burial some
seeds of 51 of the 107 species were still alive. They were primarily weed
seeds and hard-seeded legume species. Of 31 cultivated species in the
experiment, 22 were dead after 20 years in the soil and most of them
after 1 year. The following cultivated species showed some germination
after 20 years: Alsike clover, beet, bushclover, celery, Kentucky blue-
grass, red clover, timothy, tobacco, and white clover. Forty-four spe-
cies grew after 30 years and 36 species after 39 years. The experiment
was discontinued in 1941 (after 39 years) just before the Arlington
Experimental Farm of the U.S. Department of Agriculture was occu-
pied by the Defense Department.
A seed longevity experiment was started by sealing 20 lots of 120
kinds of seeds in 2,400 glass tubes (Went, 1948). The seeds were dried
before they were stored in vacuum tubes. One lot of each kind of seed
was to be removed at 10-year intervals at first and later at 20-year
intervals until A.D. 2307. The object of this experiment was to obtain
data on (1) how long seeds can be stored without losing their power to
germinate, (2) how they are affected by long storage, and (3) what
evolutionary changes occur from year to year in plants of the same
species.
Circumstantial Evidence of Long Lifespans of Seeds

A large number of presumed long lifespans of seeds can be cited. The
age of the seeds has been estimated by such comparisons as with objects
of known age, surroundings, geological events, and carbon-14 dating. In
some cases, the seeds have been tested by reliable procedures, but in
other instances, germination in nature has been observed and accepted
as fact. Observations of the appearance of seedlings cited as evidence of
seed longevity or persistence of life in seeds include the following
phenomena: Removal or burning of a building, disturbance of grassland
or forest soils, disturbance of old or ancient mounds, disturbance of the
earth's surface by bombing, drainage of old lake or pond beds, and
erosion and deposits left by glaciers. Superficially, events arising from
these phenomena appear as striking examples of seeds of great longev-
ity. However, such observations are based largely on speculation with-
out considering all the facts.
Turner (1933) pointed out the weakness of conclusions based on such
observations in the following words: "When the longevity of buried
seeds is estimated entirely from the circumstances in which they are
found, there is the possibility of error, and the only satisfactory
methods of ascertaining the length of time that buried seeds will remain
viable are by experiments such as those begun by Beal and Duvel in the
United States." Turner (1933) described 20 examples of claimed lon-
gevity of buried seeds about which the age of the seed can be ques-
tioned. We describe five examples, some of which were also described
by Turner (1933).
Seeds in Soil Disturbed by Digging Graves
In his "Flora of the Summe Battlefield," Hill (1917) gave the following
account: "In July poppies ( Papaver rhoeas L.) predominated, and the
sheet of colour as far as the eye could see was superb; a blaze of scarlet
unbroken by tree or hedgerow. Here and there long stretches of
chamomile (Matricaria chamomilla L.) broke into the prevailing red
and monopolized some acres; and large patches of yellow charlock were
also conspicuous, but in the general effect no other plants were notice-
able, though a closer inspection revealed the presence of most of the
common weeds of cultivation. . . . Charlock (Sinapis arvensis L.) not
only occurred in broad patches but was also fairly uniformly distrib-
uted, though masked by the taller poppies. Numerous small patches
were, however, conspicuous and these usually marked the more re-
cently dug graves of men buried where they had fallen. . . . No doubt
in the ordinary operations of ploughing and tilling of the ground in years
before the war much seed was buried which has been brought to the
surface by the shelling of the ground and subsequent weathering. In
this connection the presence of charlock on the more recently dug
graves, where the chalk now forms the actual surface, is of interest,
since it adds further proof of the longevity of this seed when well buried
in the soil."
Actually there is no proof that the charlock seeds (Brassica kaber
(DC.) L. C. Wheeler) (=Sinapis arvensis) had been buried either deep
or for a long time. The seeds may have been lying in the soil but close to
the surface, where conditions were not conducive to germination until
they were brought to the surface, where light and oxygen were avail-
able and where any accumulated carbon dioxide within the seeds could
escape.
Seeds in Soil Under Forests of Known Age

Peter (1893) reported studies made of the seed content of soils of a
forest that had been planted on meadows and pastures for known
periods and kept free from open land plants by shading. In general, as
the age of the forests increased, seeds of field plants became more
scarce. He found seeds of Hypericum humifusum, Juncus bufonius,
and Stellaria media in deep soil layers of forests 100 years old. In soils
of forests 20 to 46 years old he found seeds of a large number of open
land plants belonging to such genera as Anagallis, Chenopodium,
Juncus, Plantago, Polygonum, Sinapis, Stachys, Stellaria, and Thla-
spi. Peter (1893) concluded that seeds of some meadow and swamp
plants that may lie in the soil more than 50 years are still capable of
germination.
His conclusions appear to be correct, and in lieu of a planned ex-
periment they provide some worthwhile data; however, his data would
be more valuable if there were no question about the age of the seeds.
Seeds Frozen in Arctic Tundra

Porsild et al. (1967) reported that seeds of arctic lupine, at least
10,000 years old, found in lemming burrows deeply buried in perma-
nently frozen silt of the Pleistocene age, germinated and grew into
normal, healthy plants. A mining engineer found the seeds in the
permanently frozen burrows associated with the remains of a nest, fecal
matter, skulls, and skeletons of a rodent, later identified as the collared
lemming. According to Porsild et al. (1967), the mining engineer kept
the skeletons in a dry place for 12 years, when the authors learned of
the specimens and historical background.
Through carbon-14 dating, comparison with other archeological
artifacts, and natural and geological history of the central Yukon area
where the seeds were found, they concluded that the seeds had been
placed in the burrows by the lemmings and were at least 10,000 years
old. About two dozen seeds easily identified as those of the arctic lupine
were found, and about half were remarkably well preserved. Some of
the best preserved seeds yielded six seedlings when tested for
germination and eventually yielded normal, mature plants, indistin-
guishable from those grown from fresh seeds. The collared lemming no
longer occurs in the central Yukon, but the arctic lupine is a common
species of tundra and subalpine forests of northern and central Alaska,
the Yukon Territory, and the northern Mackenzie District.
A critical review of this account reveals the weaknesses of evidence
supporting the claimed age of these seeds.
Canna Seeds in Burial Mound
Researchers in Argentina (Sivori et al., 1968) reported that three
seeds of a Canna species (exact species not determined) were obtained
from a tomb being excavated in Santa Rose de Tastil. The seeds were
inside walnut shells strung in a necklace to form rattles. As determined
by carbon-14 dating tests of cameloid bones from overlying strata of the
site, the seeds were about 530 years old. This information, supple-
mented by the fact that no Incaic elements were found in the burial
mound, suggests that the city was developed before 1420. Two of the
three seeds were tested for germination and subsequent growth. The
first seed germinated, but the radicle ceased to grow, even after adding
gibberellic acid and indole-3-acetic acid. Another seed was tested under
aseptic conditions on a medium containing minerals and growth regula-
tors. It germinated and continued growth after transplantation into
quartz gravel with a nutrient solution in a greenhouse. The roots of the
plant exhibited irregular geotropic response, even after being trans-
planted in the greenhouse.
The principal weakness in this account of seed longevity is the
accuracy of the carbon-14 dating test and the time relationship of the
cameloid bones tested and the Canna seeds. (See carbon-14 dating
results by Libby (1951) and Godwin and Willis (1964)for Nelumbo seeds
in the next section.)
Indian Lotus Seeds Buried in Former Lakebed
Probably the longest claimed longevity for any seed is that recorded
by Ohga (1923). He obtained approximately 100-percent germination
with seeds of Indian lotus, which he found in a peat bed buried 2 feet
deep with loess in the Pulantien River valley in southern Manchuria.
The bed was 41 feet above the present water level of the river. Judging
from the age of the weeping willows on the bed of this former lake and
from the lowering rate of the water level, Ohga (1923) concluded that
the seeds were probably at least 120 years old. Possibly they were 400
years old or even older judging by the rate of erosion. Although the
supply of seeds was limited, Ohga deposited some with the Tohuku
Imperial University in Japan, and in 1926 he gave 30 seeds to the
British Museum in London.
Later Chaney (1951) obtained a few seeds from the Ohga collection
and made them available for study in the United States. Records are
available (Wester, 1973) showing that in tests made in Japan, Eng-
land, and the United States on 156 seeds all germinated except 2. A dry
seed from the Sloane collection in the British Museum germinated after
237 years (table 30). Two seeds from the Ohga collection, tested for
germination in 1951 by Wester, are illustrated in figure 33.
The age of these Nelumbo seeds is perhaps the most controversial
subject in the field of seed longevity with a legitimate basis. The
discoverer of the original seedbed and seeds estimated the seeds to be
at least 120 years old and perhaps as old as 400 years (Ohga, 1923).
Later Chaney (1951), a paleontologist, reevaluated the information on
the age of the seedbed and concluded that the seeds might be as much as
50,000 years old.
Arnold and Libby (1951) reported that they had determined by
carbon-14 dating that the seeds were 1,040 ± 210 years of age. This age
was generally accepted until Godwin and Willis (1964) conducted car-
bon-14 dating tests and concluded the correct age to be 100 ± 60 years.
Wester (1973) reviewed the entire history and arrived at an age of
1,024 years, which agrees with Libby's findings of 1,040 ± 210 years.
There is little question that these seeds are old, but their age has not
been established to the satisfaction of biologists. Like seeds of many
leguminous species, the seeds of the Indian lotus plant have very hard
seedcoats, which resist decay and are impervious to water and gases.
Also, the peat in which the seeds were found might have retarded the
deterioration of the seedcoat.
Life in Mummified Seeds

The following article on "Wheat 2,000 Years Old" appeared in the
Sunday magazine section of a Washington, D.C., daily newspaper,
dated January 13, 1957:
"This story begins more than 2,000 years ago, when relatives of a
deceased Egyptian nobleman buried some wheat kernels in his tomb,
presumably so the departed could grow his own in the land of the dead.
"The nobleman was hardly in any shape for showing as a museum
mummy when unearthed recently by a Swiss archeologist, but the wheat
seeds appeared to be in good condition. The archeologist, no farmer
himself, wondered if they might still sprout under proper cultivation.
"Reviewing his proposal to test the grain under various conditions of
soil and clime, the Egyptian government turned thumbs down on
experimenting anywhere but in Egypt. But placing agronomy ahead of
sovereignty, the scientist whisked a handful of the kernels out of the
country.
FIGURE 33. —Seeds of Nelumbo nuctfera alleged to be over 1,000 years old: Above, before
absorbing water; below, after germination.
"Distributing it to several experimental farmers in Europe, he seems

to have scored a major success with the wheat only in Sweden. There,
Oscar Johnson of Smaland planted three kernels, carefully nursed them
through the season and ended up with several thousand seeds for the
next planting. By present:, count, he has enough to sow 2 1/2 acres. Should
he have a successful harvest next year, he expects to reap enough grain
to plant a few hundred acres. From then on, the wheat that had laid
dormant for ages may be available for others.
"Called "Osiris," the reborn wheat strain has thus far proved to have
tremendous hardiness. What other beneficial qualities it may have, Mr.
Johnson and other agriculturists the world over are interested in
finding out."
Two photographs accompanied the article. The caption to one photo-
graph showing four spikes of branched grain reads as follows: "Buried
for centuries in an Egyptian tomb, wheat kernels were unearthed and
sown by Oscar John Johnson of Sweden, who produced this initial
yield." The other photograph, showing a man who apparently is trans-
planting seedlings of grain in soil, has the following caption: "Farmer
Johnson had only three grains of the 2,000-year-old wheat, so he planted
them in a special bed where he could keep a watchful eye on their
progress. All three grains prospered in spite of their age. The matured
wheat exhibited extreme hardiness, and he reaped enough to plant
more than two acres now."
This article was sent to the head of the Swedish Seed Testing Station
at Solna, Sweden, requesting that he comment on the article. Following
is his reply, dated February 1957:
"This is the same old story that appears in the world press at least
once in 10 years. The Egyptians by now know very well that it is a lie. If
living wheat kernels are found in ancient tombs, they were pr there
rather recently by mice or other animals. Cereals deposited there 2,000
years ago have now turned to pure coal and are as dead as they can be.
When staying in Egypt 10 years ago I had an opportunity to convince
myself personally about this. Since no business can be done with such
seeds in Egypt, the kernels are sent to other countries and are sold at
fancy prices.
"The special case of Mr. Johnson first circulated in the Swedish press
a few years ago and then has been taken up by newspapers and journals
in most other countries. About a year ago I had a letter from Canada
asking my opinion of it. It is a pity that journalists always copy
sensational stories but never disprovals of them! Mr. Johnson's wheat is
a variety grown today in Egypt."
Other similar stories of mummy seeds are discussed by Brooke
(1935), Luthra (1936), and White (1946).
A form of wheat sold to tourists as "miracle wheat" or "mummy
wheat" is a common branched form of Triticum turgidum, which grows

in southern Europe and northern Africa. It is commonly cultivated as a
curiosity (Turner, 1933). Similarly, fasciated forms of a pea similar to
the "mummy pea" were illustrated in a European herbal as early as
1590. One form of the pea was described and named Pisum umbellatum
(rose or crown pea) in 1771 (White, 1946).
Turner a933) of Kew Gardens, London, indicated that there was no
authenticated evidence that wheat taken from undisturbed Egyptian
tombs will germinate. An experiment was made at Kew Gardens with
grain from a model granary found in a tomb of the 19th dynasty.
Samples were tested under various conditions. The effect of colored
glass was tried to induce germination, but after 3 months the grain had
turned to dust.
We have confirmed these results in tests made at Beltsville, Md.,
using mummified grain from the Anatolian excavations at the site of
Alishar. Information from the Oriental Institute, University of Chi-
cago, indicates that the material dates back to at least 700 to 1200 B.C.
and possibly to about 3000 B.C.
FIGURE 34.—Seeds of 1973 barley and wheat crops compared with carbonized structures
("seeds") of the same species from the Alishar excavations, dating from approxi-
mately 700 to 3,000 B.C.: Top to bottom, barley from Alishar and 1973 crop; wheat
from Alishar and 1973 crop.
The printed word does not seem to dispel the story of life in mummy
seeds as such stories appear in the popular press from time to time. For
this reason, we are showing some of the seeds from the Alishar
excavations along with seeds harvested in 1973 (fig. 34). The so-called
mummy seeds have retained the shape of barley and wheat, but the
structure is similar to that of charcoal. There is no possibility of these
structures producing seedlings. Turner (1933) said, "It is popularly
asserted that miracle wheat and mummy peas originate from Egyptian
tombs and that such seeds germinate when sown, but in every instance
the statements prove to be without foundation."
GLOSSARY
Absolute humidity.—Amount of water vapor actually in the air, ex-
pressed either in its expansive force or in its weight per given
volume, as grains per cubic foot.
Absorption. —Imbibing of water by living cells or tissues in a seed.
Accelerated aging test.—Intentionally subjecting seeds to adverse
storage conditions for a short period to estimate their possible life-
span under favorable storage conditions.
Achene.—Small, dry, one -seeded fruit with thin distinct wall, which
does not split open.
Actinomycetes.—Bacterialike micro-organisms with ribbonlike form.
Activated alumina.—Highly porous, granular form of aluminum oxide,
with high adsorptive capacity for moisture.
Adsorption.—Taking up a gas, vapor, or dissolved material on the
surface of a solid; finely divided materials, as active carbon and silica
gel, can adsorb relatively large quantities of other materials.
Aeration.—Movement of outside air through stored seeds to prevent
heating and to facilitate drying.
Aging test.—Any test to determine the deterioration of seeds with their
increased age, usually a germination or vigor test.
Air-dry.—Sufficiently dry so that no further moisture is given off on
exposure to air; i.e., in moisture equilibrium with the surrounding air.
Air-oven method.—Drying seeds in a forced-air oven to determine their
moisture content.
Alpha amylase.—An enzyme occurring widely in seeds, leaves, and
other plant parts; it accelerates starch hydrolysis.
Ambient.—That which encompasses or surrounds on all sides a desig-
nated object, state of matter, system, etc.; as used here, ambient
refers to temperature or relative humidity of the atmosphere in the
natural state without modification by man.
Anabolic process.--Process by which matter is changed into tissues of
living plants or seeds.
Axillary bud.—Bud developed in the angle between leaf and stem.

Backcrossing.—Crossing a hybrid with one of the original parents.
Bacteria.—Simple microscopic vegetable organisms usually single-
celled and without chlorophyll, multiplying by fission (splitting apart)
and spore formation.
"Baldheads."—Bean seedlings with the primary leaves missing and the
terminal bud present or absent.
Batch drying.—Drying seeds in relatively small quantities while sta-
tionary as opposed to drying in a continuous moving line.
Biochemical change.—Change in the chemistry of a plant, plant part, or
seed.
Bracts.—More or less modified leaves subtending a flower or belonging
to an inflorescence.
Bruchids.—Family of small vegetarian beetles; the larvae live mainly in
the seeds of leguminous plants.
Bunt spores.—Reproductive units of certain fungi that attack cereal
plants.
Caryopsis.—One-seeded fruit with pericarp and seedcoat fused into one
covering, as in corn and other grains and grasses.
Catabolic process.—Pertaining to or characterized by the release of
energy.
Chaff.—Glumes or husks of grains and grasses separated from the seed
by threshing, winnowing, etc.
Chalcids.—Minute wasplike insects belonging to the order Hymenop-
tera, a few species of which invade seeds.
Check samples.—Samples of original seed lots used as a basis for
establishing the effects of any special treatment on the response of
seeds of the same lots. For example, seeds held at room temperature
constitute the check sample for seeds of the same lot stored at a high
temperature and high relative humidity and at a low temperature and
low relative humidity.
Chromosome aberrations.—Deviations from normal chromosome
structure.
Chromosome 10.—Geneticists have assigned numbers to various chro-
mosomes; 10 identifies a specific chromosome.
Closed system.—Completely enclosed, sealed system; device, con-
tainer, or combination of devices designed to perform a specific
function within a sealed environment, as the refrigeration system for
a domestic refrigerator.
"Cold Test. "—Special germination test; for corn, seeds are placed in a
very moist sand/soil/peat moss mixture, then held at 10° C for 7 days
before being placed in the regular germination temperature.
Coleoptile.—Sheathlike leaf of grasses and other monocotyledons; it
protects the delicate growing point as it emerges from the soil.
Coleorhiza.—Sheath surrounding the radicle in some plants, through

which the root bursts.
Combine thresher.—Machine that cuts, threshes, and cleans grain while
moving over the field.
Convection.—A type of air current.
Cotyledons.—Seed leaves of the embryo, usually thickened for storage
of reserve food; they may serve as true foliage leaves.
Cultivar.—Cultural or cultivated variety.
Culture medium.—Laboratory substance on which micro-organisms
and excised embryos may be grown.
Cuticle.—Very thin detachable skin covering a plant, especially the
leaves.
Cytological change.—Change in the intrinsic character or function of a
cell.
Decorticated seed.—Seed with its outer covering removed.
Dehumidification.—Process by which the water is removed from a
substance; moisture vapor is removed from air.
Dehumidifier.—Equipment used to remove humidity (moisture) from
air.
Desiccant.—Drying agent.
Desiccator.—Usually a glass jar fitted with an airtight cover and
containing at the bottom some desiccating agent, such as calcium
chloride.
Desorption.—Removal of a substance from an absorbed state.
Dielectric properties.—Properties that make a material a nonconductor
of electricity.
Dough stage.—Stage in development of cereal grains when the interior
of the kernel (seed) is of doughlike consistency.
Dry-bulb thermometer.—Ordinary thermometer with its mercury-con-
taining bulb kept dry. (See psychrometer.)
Dry heat.-----Heat applied by dry air as opposed to moist heat, which is
applied by moist air.
Dry weight.—Moisture-free weight.
Dynamic method.—Procedure for determining equilibrium moisture
content by bubbling air through absorption towers containing an acid
or saturated salt solution, which controls the humidity around seeds
or other material.
Economic plants.—Plants of practical utility; commercial plants.
Ecosystem.—System involving all ecological or environmental factors,
such as climate, soil, physiography, and associated organisms.
Electric hygrometer.—Electric instrument for measuring the degree of
moisture in the atmosphere.
Electric moisture meter.—Electrical instrument that measures the
moisture content of seeds and other materials.
Embryo.—Rudimentary plant within the seed.

Embryo axis.—Hypocotyl-root axis bearing at one end the root
meristem and at the other the cotyledon or cotyledons and the shoot
meristem.
Embryo transplant technique.—Procedure by which the embryo of one
seed is transplanted to the endosperm of another seed.
Endogeocarpic flora.—Certain microscopic plants that live or are found
in the soil and can invade plants.
Endosperm.—Tissue of seeds, developing from fertilization of the polar
nuclei of the ovule by the second male nucleus; it nourishes the
embryo.
Enzyme.—Catalyst produced in living matter; a specialized protein
capable of aiding in bringing about chemical changes; promotes a
reaction without itself being changed or destroyed.
Ergot.—Hard resting body produced by the fungus Claviceps pur-
purea; also the name of a disease of rye and other grasses.
Field emergence.—Emergence of seedlings from seeds planted in the
field; usually refers to percentage of seeds planted in the field that
germinates and produces seedlings strong enough to emerge through
the soil.
Field fungi.—Molds that live outside on growing crops.
Food transport system (in embryo).—System by which food is moved
from storage tissues of the seed to growing tips of the root and shoot.
Forage.—Vegetable food of any kind for animals, especially that con-
sumed by domestic animals as pasture or browse.
Freeze-drying.—Process for drying seeds and other materials; it uses a
freezing temperature combined with moving dry air, which sublimes
frozen water to water vapor.
Fruit(s).—Ripened (matured) ovary of a_ plant, together with any in-
timately attached parts that developed with it from the flower. Fruits
of buckwheat, cereals, grasses, sunflower, and many others are
commonly called seeds.
Fumigant.—Chemical applied as a gas to destroy molds, insects, ro-
dents, etc.
Fungi.—Lower plant forms devoid of chlorophyll and comprising
molds, mildews, rusts, smuts, mushrooms, toadstools, puffballs, and
allied forms that reproduce by spores.
Fungicide.—Chemical used to kill fungi.
Gall(s).—Swollen plant part caused by presence or action of an insect or
nematode.
Gaseous water.—Water vapor.
Gas storage.—Storage in an atmosphere composed of a gas other than
air.
Genera.—Plural of genus; groups of structurally or phylogenetically

related species; categories of classification ranking between family
and species.
Genetic stock.—Strain of seeds used in studies pertaining to the genesis
of a particular plant; its mode of production and development; strain
used for genetic studies.
Genotype.—Hereditary makeup of an individual plant, which, with the
environment, controls the individual's characteristics, such as type of
flower or shape of leaf.
Germinate.—Initiate growth or development, especially when refer-
ring to a spore or seed.
Germ plasm.—Living substance of the cell nucleus that determines the
hereditary properties of organisms and that transmits these proper-
ties from makeup of organisms. For example, geneticists and plant
breeders often refer to the seeds and plants used in their research
and breeding as their "collection of germ plasm."
Glumes.—Chafflike bracts; specifically, two empty chaffy bracts at the
base of spikelets in grasses.
Grain.—Seed or seedlike fruit of any cereal grass, such as millet, oats,
rice, and wheat.
Grain head.—Attachment for a combine designed specifically for har-
vesting grain.
Hair hygrometer.—Instrument in which expansion and contraction of a
hair is used to measure changes in the amount of moisture in the air.
Hard seed.—Seed with seedcoat impervious to water or oxygen re-
quired for germination, or to carbon dioxide that may accumulate
within the seed and retard or prevent germination; sometimes over-
come by scratching or scarifying the seedcoat or removal by brief
immersion in concentrated sulfuric acid and thorough washing.
Hectare.-1 ha is 2.4710 acres.
Herbarium specimen.—Dried and pressed specimens of plants mounted
or otherwise prepared for permanent preservation and systematic
arrangement in a room, building, or institution. (herbarium) where
such collections are kept.
Hermetically sealed.—Sealed so that no gas or moisture can enter or
escape.
Hermetic storage.—Storage in an airtight container.
Hilum. —Scar or point of attachment of the seed.
"Hot spots. "—Areas in a mass of stored seeds or grain where tempera-
ture is as high as 660 C or higher.
Hybrid.—Offspring of the union of a male of one race, variety, species,
genus, etc., with a female of another; a crossbred plant.
Hydrophobic.—Lacking a strong affinity for water.
Hygroscopic.—Readily absorbing and retaining moisture, as caustic

potash, or becoming coated with water, as glass.
Hygroscopic equilibrium measurement.—Measuring the amount of
water in or on a substance when there is no movement of water
between the substance and the surrounding air.
Hygroscopicity.—Denotes sensitivity to moisture.
Hygroscopic moisture absorption.—Absorption of moisture from the
surrounding air.
Hygrothermograph. —instrument that records both humidity and tem-
perature on the same chart.
Hysteresis effect. —Phenomenon when at a given relative humidity
seeds or rain may reach two different equilibrium moisture con-
tents- one by increasing the relative humidity from a low level and
another by decreasing the relative humidity from a high level.
Illumination.—Supplying light.
Immature seed.—Seed not fully developed.
Inbred.—Successively self-fertilized; also, a plant or progeny resulting
from successive self-fertilization.
Inert. —Lifeless.
Interstitial air spaces.—Air spaces between the cells in a plant tissue or
inside a seed.
Layer drying.—Drying seeds in thin layers as opposed to a large mass
of seeds.
Leguminous plants or legumes.—Members of the pea or legume plant
family (Leguminosae).
Lemma.—Small greenish bract that is part of the floret in grasses.
Lifespan. —Length of time a particular lot or kind of seeds will survive
under favorable storage conditions.
Lipids.—Group of substances comprising the fats and other esters that
possess analogous properties.
Lipolytic degradation.—Decomposition of fat.
Longevity.—Long duration of life, length of life, or lifespan.
Luteus genes.—Genes that carry a yellow plant color factor; they are
lethal as they prevent normal chlorophyll development.
Macromolecular water layer.—Water layer surrounding a macromole-
cule.
Macromolecule.—Large molecule; grouping together of simple mole-
cules capable of independent existence as in cellulose.
Malt agar.—Culture medium used for laboratory cultivation of certain
fungi.
Malvaceous plant.—Plant belonging to the mallow family, as cotton and
okra.
Manifold.—Chamber from which air is distributed to more than one
duct.
PRINCIPLES AND PRACTICES GF SEED STORAGE
Mature stage.—Completely developed; full grown.

Membrane.—Any thin, soft, pliable sheet or layer of animal or vegeta-
ble origin; a limiting protoplasmic surface.
Mercurial.—Chemical compound containing mercury.
Metabolism.—Chemical changes occurring in an organic body in living
and growing processes.
Metabolites.—Products of metabolism.
Microflora.—Microscopic plants as a whole that inhabit a given area,
such as a seed.
Micropyle. —Pore or opening through which the pollen tube enters the
embryo sac during fertilization and through which the radicle fre-
quently emerges when the seed germinates.
Milk stage.—Stage in the development of the kernel of a grain when its
contents are a milklike liquid.
Moisture-barrier material.—Any material through which moisture ei-
ther cannot move or moves very slowly.
Moisture content equilibrium.—State when the moisture content in the
air and in the seed remains constant; no movement of moisture either
into or out of the seed.
Moisture equilibrium.—Relative humidity of the air is in equilibrium
with the moisture content of the seed.
Moisture gradient.—Difference in moisture concentration inside and
outside a seed and between the seed surface and the air surrounding
the seed.
Moisture translocation.—Movement of moisture from one part of a
seed to another.
Multwall bag.—Bag with multiple walls, usually two or more, com-
posed of paper, foil, plastic film, or cloth, alone or in various combi-
nations.
Mutagenic changes.—Change in the genetics of a seed; change in form
or qualities of chromosomes.
Mycoftora.—Fungal component of a microflora.
Nematode.—Microscopic wormlike animal.
Nomograph.—Alinement chart; graph that enables one, by the aid of a
straight edge, to read the value of a dependent variable when the
value of the independent variable is given.
Nutlet.—Any small nutlike fruit or seed, as in the mint and vervain
plant families.
Oilseed.—Any seed from which oil is expressed; a seed containing a
large percentage of oil in its storage tissues.
"Once-over harvesting. "—Harvesting the entire crop at one time, as
opposed to going over a field several times to complete the harvest.
Open storage.—Storage with free access to normal atmospheric condi-
tions.
Ordinary storage.—Storage as usually practiced with no attempt to

control either temperature or relative humidity of the storage area.
Osmophilic.—Preferring to grow with very little water.
Overwinter storage.—Storage through the winter from harvest in the
fall to planting time in the spring.
Ovules.—Bodies within the ovary of a flower that become seeds after
fertilization and development.
Oxidation.—Act or process of being oxidized.
Palea.—Upper bract that with the lemma encloses the flower in
grasses. (See glumes.)
Pallet box.—Box in which seeds are placed for handling during proc-
essing and that is moved by forklift. (See totebox.)
Parasitic fungi.—Molds that exist at the expense of other organisms.
Percent germination.—Percentage of seeds that produces normal
seedlings when tested in a laboratory according to established pro-
cedures.
Pericarp.—Seed covering derived from the ovary wall; it may be thin
and intimately attached to the seedcoat as in a kernel of corn, fleshy
as in berries, or hard and dry as in pods and capsules.
Photoperiod.—Length of the light period on a given day.
Photoperiodism.—Response to the timing of light and darkness.
Photosynthetic organs.—Organs containing chlorophyll and capable of
photosynthesis.
Physiological dormancy.—Dormancy caused by disorganization of
functions or of metabolism as brought about by extremes of temper-
ature, oversupply of water, etc.
Placenta.—Any sporangia-bearing surface; in seed plants that part of
the carpel bearing ovules.
Planting stock.—Seeds selected and preserved especially for planting,
as opposed to seeds held for human or animal food.
Plumule.—Major young bud of the embryo within a seed or seedling
from which the aerial part of the plant will develop.
Premilk stage.—Stage in the development of a seed that precedes the
milk stage.
Primary root.—First root of a plant that develops from the radicle.
Propagation.—Perpetuating and increasing plants vegetatively.
Protoplasm.—Essential, complex, living substance of cells on which all
the vital functions of nutrition, secretion, growth, and reproduction
depend.
Provenance.—Origin, source, place where found.
Psychrometer.—Hygrometer or instrument for measuring the water
vapor in the atmosphere, consisting essentially of two similar ther-
mometers, the bulb of one kept dry, the other wet. The wet bulb is
cooled as a result of evaporation and consequently shows a tempera-
ture lower than that of the dry-bulb thermometer. Since evaporation

is less on a moist day, the difference between the thermometer
readings is greater when the air is dry. This difference constitutes a
measure of the dryness of the surrounding air.
Psychrometric chart.—Graphical representation of psychrometric data,
showing the relationship between dry-bulb temperature, dewpoint,
wet-bulb temperature, and relative humidity.
Radicle.—Rudimentary root or lower end of the hypocotyl of the
embryo; it forms the primary root of the young plant.
Relative humidity.—Ratio of the quantity of water vapor actually in
the air to the greatest amount possible at a given temperature.
Respiration.—Oxidation-reduction process occurring in all living cells
whereby chemical action occurs, producing compounds and releasing
energy that are partly used in various life processes.
Respiration quotient (RQ).—Ratio of the volume of carbon dioxide
(CO 2) given off in respiration to that of the oxygen (0 2 ) consumed-
RQ = CO 2/02 .
Root axis.—Principal line along which a root may grow; hypothetical
central line of a root.
Sample.—Part of a seed lot presented for inspection or shown as
evidence of the quality of the whole lot.
Saprophytic fungi.—Molds that live off nonliving material.
Scarified seed.—Seed that has had its coat scratched to permit ready
entry of water and exchange of gases.
Schematic flow.—Movement of seeds according to a previously de-
signed scheme or pattern.
Sclerotia.—Hard, resting bodies produced by certain fungi.
Scutellum.—Specially differentiated cotyledon in grass seeds; shield-
shaped organ through which the embryo absorbs food from the
endosperm.
Seated storage.—Storage in a sealed container; usually refers to her-
metic storage.
Secondary root.—Lateral branch of the primary root.
Seed(s).—Mature ovule consisting of an embryonic plant together with
stored food, all surrounded by a protective coat.
Seedcoat.—Outermost tissues or "skin" of a seed; sometimes this coat is
extremely hard and waterproof, preventing entrance of water to
initiate germination unless it is broken, scratched, or eroded.
Seedling.—Young plant grown from a seed.
Seed lot.--Specific quantity of seed usually assigned an identifying
number or symbol; examples are the seed from a single field or a day's
harvest.
Seed phyiology.—Branch of biology dealing with the organic processes
and phenomena collectively of seeds and their parts.
Seed stock.—Quantity of seed reserved for planting for the production

of a new seed crop.
Shoot axis.—Principal line along which a shoot may grow; hypothetical
central line of a shoot.
Sigmoid.—Curved in two directions, like the letter "S."
Silica gel.—Regenerable adsorbent consisting of amorphous silica.
Smut ball.—Reproductive body produced by smut fungi.
Species.—Group of closely related organisms; for example, Medicago
saliva is the botanical name for alfalfa. Medicago is the genus and
saliva is the species. Several species belong to the genus Medicago;
for example, M. lupulina (black medic) and M. orbicularis (button-
clover).
Spine(s).—Any stiff sharp process distinguished from a thorn by the
absence of vascular tissue. Spines are frequently found on leaf
margins as in thistles.
Stand of plants.—Relative number or distribution of plants growing
over a given area, especially soon after germination.
Static efficiency.—Efficiency of static pressure in moving air through a
layer of seeds.
Static method.—Method by which a large fan builds up air pressure
against a mass of seeds in a dryer in order to move more air through
the seeds so as to speed up the drying process.
Static pressure.—Pressure acting by mere weight (force) without mo-
tion; pressure exerted by mere mass at rest.
Storage fungi.—Fungi that live off stored seeds and grains.
Stress test.—Test in which seeds are intentionally subjected to the
action of adverse external forces, usually unfavorable germination
conditions.
Strophiole.—Swollen appendage at the hilum of certain seeds, as that of
spurge.
Substrate.—Substratum, a substance acted upon.
Substratum.—Seedbed in a germination test; soil is a substratum for
germinating seeds.
Temperature gradient.—Rate of temperature change with increase in
height; lapse rate or change with distance from heat or cooling source.
Terminal bud.—Bud growing at the end of a branch or stem.
Testa.—Hard external coating or integument of a seed; seedcoat.
Thermocouple.—Thermoelectric couple used to measure temperature
differences.
Threshed.—Beat out from the head, pod, or capsule; separated from the
plant.
Totebox.—Open or closed metal or wood box, which is placed on a pallet

and is moved by a forklift. (See pallet box.)
Treated seed.—Seed to which a poisonous chemical compound has been
applied to kill pests.
Umbel(s).—Inflorescence in which the peduncles or pedicels of a cluster
spring from the same point, as in carrot, coriander, or parsnip.
Unthreshed.—Not beat out; not separated from the pod, head, or
capsule.
Vacuum storage.—Storage in a sealed container from which all or
nearly all air was evacuated before sealing; storage in a (partial)
vacuum.
Vascular.—Furnished with vessels or ducts.
Viability.—Quality as state of being alive; ability to live, grow, and
develop, as the viability of certain grains under dry conditions.
Viability test.—Test to determine whether a seed or the percentage of
seeds in a lot is alive or dead.
Viable.—Capable of growing or developing, as viable seeds.
Vigor.—Condition of active good health and natural robustness; when
seeds are planted, vigor permits germination to proceed rapidly and
to completion under a wide range of conditions.
Virus.—Extremey small, filterable body, which multiplies only in
living cells.
Vitality.—Germinating power.
Water-curtain germinator.—Germinator (germination chamber) that is
heated, cooled, aerated, and humidified by means of a curtain of
water falling down the back of the chamber; water is recirculated.
Weevits.—Snout beetles of the suborder Rhynchophora with snout-
bearing jaws at the tip that damage stored seeds and grain.
Wet-bulb thermometer.—Ordinary thermometer with mercury-con-
taining bulb that is kept wet. (See psychrometer.)
Wet weight.—Weight before drying.
Windrow.—Row of hay raked up to dry before being stacked or baled;
also any similar row for drying, as of grain or peanuts.
Xerophyte.—Plant structurally adapted for growth with a limited water
supply.
Yellow cuprocide.—Copper-containing fungicide used to protect seeds
from soil fungi.
Yield.—Aggregate of products resulting from growth or cultivation;
quantity produced, such as 100 bushels of corn per acre.
Zygote.—Fertilized egg, potentially capable of developing into a seed.
VERNACULAR AND BOTANICAL-ZOOLOGICAL

NAME EQUIVALENTS 11
Plants
Achillea, the pearl—Achillea ptarmica L.
Ageratum—Ageratum mexicanum Sims
Alfalfa—Medicago saliva L.
Alyceclover—Alysicarpus vaginalis (L.) DC.
Alyssum—Alyssum maritimum (L.) Lam.
Amaranth, redroot—Amaran thus retroflexus L.*
Amaranthus—Amaran thus spp.
Anemone—Anemone spp.
Angel-trumpet—Datura arborea L.
Anise—Pimpinella anisum L.
Apple—Malus sylvestris Mill.
Arabis--Arabis atpina L.
Armeria—Armeria spp.
Artichoke—Cynara scolymus L.
Asparagus—Asparagus officinalis L.
Asparagus, fern—Asparagus plumosus Baker and A. sprengeri Regel
Asparagusbean, sitao— Vigna unguiculata subsp. sesquipedalis (L.)
Verde.
Aster—Aster spp.
Aster, China—Caltistephus chinensis (L.) Nees
Babysbreath—Gypsophila spp.
Bachelor's button, cornflower—Centaurea cyanus L.
Bahiagrass--Paspalum notatum Fluegge
Balloonflower—Platycodon grandiflorum DC.
Balm—Melissa officinalis L.
Balsam—Impatiens spp.
Barley—Hordeum vulgare L.
Barnyardgrass, junglerice—Echinochloa spp.*
Basil, sweet—Ocimum basilicum L.
Basketflower—Centaurea americana Nutt. and Hymenocallis cala-
thina Nichols
Bean:
Field—Phaseolus vulgaris L.
Garden (snap)—Phaseolus vulgaris L.
Kidney—Phaseolus vulgaris L.
Lima—Phaseolus vulgaris L.
Mung— Vigna radiata (L.) Wilczek (syn. Phaseolus aureus Roxb.)
Bean—Continued
Navy—Phaseolus vulgaris L.
Scotch— Vicia faba L.
Tapilan— Vigna umbeliata (Thunb.) Ohwi and Ohashi (syn. Pha-
seolus calcaratus Roxb.)
Beet:
Field—Beta vulgaris L.
Garden—Beta vulgaris L.
Sugar—Beta vulgaris L.
Beggarweed—Desmodium tortuosum (Sw.) DC.
Begonia—Begonia spp.
Bellflower:
Bluebell—Campanula spp *
Peach—Campanula persicifotia L.
Bentgrass:
Colonial (including Astoria and Highland)—Agrostis tennis Sibth.
Creeping—Agrostis palustris Huds.
Velvet—Agrostis canina L.
Bermudagrass—Cynodon dactylon (L.) Pers.
Betony—Stachys spp.*
Bindweed—Convolvulus spp.*
Bluegrass:
Bulbous—Poa butbosa L.
Canada—Poa compressa L.
Kentucky—Poa pratensis L.
Nevada—Poa nevadensis Vasey ex Scribn.
Rough—Poa trivialis L.
Texas—Poa arachnifera Torr.
Wood—Poa nemoralis L.
Bluestem:
Big—Andropogon gerardii Vitm.
Little—Andropogon scoparius Michx.
Sand—A ndropogon hallii Hack.
Borage—Borago officinalis L.
Broadbean or horsebean— Vicia faba L.
Broccoli—Brassica oleracea var. botrytis L.
Bromegrass:
Mountain—Bromus marginatus Nees ex Steud.
Smooth—Bromus inermis Leyss.
Browallia—Browallia spp.
Brussels sprouts—Brassica oleracea vat. gemmifera DC.
Buckwheat—Fagopyrum esculentum Moench
Buffalograss—Buchloe dactyloides (Nutt.) Engelm.
Buffelgrass—Cenchrus ciliaris L. (syn. Pennisetum citiare (L.) Link)
Bugloss—Anchusa capensis Thunb.

Burclover:
California—Medicago polymorpha L.
Spotted—Medicago arabica (L.) Huds.
Burdock—Arctium lappa L. and A. minus (Hill) Bernh.
Bushclover—Lespedeza violacea (L.) Pers,
Butterflyflower—Schizanthus wisetonensis Hort.
Buttonclover—Medicago orbicularis (L.) Bartal.
Cabbage—Brassica oleracea var. capitata L.
Cabbage, Chinese—Brassica pekinensis (Lour.) Rupr.
Calceolaria—Cal ceolaria spp.
Calendula—Calendula officinalis L.
Calliopsis, dwarf and tall—Calliopsis spp.
Canarygrass—Phalaris canariensis L.
Canarygrass, reed—Phalaris arundinacea L.
Candlenut—Aleurites moluccana (L.) Wind.*
Candytuft:
Annual—Iberis umbellata L.
Perennial—Iberis gibraltarica L. and I. sempervirens L.
Canna—Canna spp.
Cantaloup—Cucumis melo L.
Canterbury-bells—Campanula medium L.
Caraway—Carum carvi L.
Cardoon—Cynara cardunculus L.
Carnation—Dianthus caryophyllus L.
Carpetgrass—Axonopus affinis Chase
Carrot—Daucus carota L.
Castorbean, Cambodia—Ricinus cambodgensis Benary*
Cathedral bells--Cobaea scandens Cav.
Catnip nepeta—Nepeta spp.
Cauliflower—Brassica oleracea var. botrytis L.
Celeriac—Apium graveolens var. rapaceum (Mill.) Gaud.
Celery—Apium graveolens var. dulce (Mill.) Pers.
Centaurea:
Royal--Centaurea. imperial i3 Hort.
Velvet—Centaurea gymnocarpa Moris and Denot
Chamomile, German (sweet false)—Matricaria chamomilla L.
Chard, Swiss—Beta vulgaris var. cicla L.
Charlock—Sinapis arvensis L. = (Brassica kaber (DC) L. C. (Wheeler))
Chervil—A nthriscus cerefolium (L.) Hoffm.
Chickweed, common—Stellaria media (L.) Vill.*
Chicory—Cichorium intybus L.
Chives—Allium schoenoprasum L.
Chrysanthemum, an —Chrysanthemum spp.
Cinchona—Cinchona ledgeriana Moens ex Trim. *

Cineraria, common—Senecio cruentus (L'Her.) DC.
Cleome, spiderflower—Cteome gigantea Hort.
Clover:
Alsike—Trifolium hybridism L.
Berseem—Trifolium alexandrinum L.
Crimson—Trifolium incarnatum L.
Ladino—Trifolium repens L.
Lappa—Trifolium lappaceum L.
Large hop—Trifolium procumbens L.
Persian—Trifolium resupinatum L.
Red—Trifolium pretense L.
Rose—Trifolium hirtum All.
Strawberry— Trifotium fragiferum L.
Striate—Trifolium striatum L.*
Subterraneum—Trifolium subterraneum L.
Suckling (small hop)—Trifolium dubium Sibth.
White—Trifolium repens L.
Cockscomb—Celosa spp.
Cocksfoot (orchardgrass)—Dactylis glomerate L.
Cockvine, thunbergia—Thunbergia alata Bojer
Cocoa—Theobroma cacao L.
Coleus, common—Coleus blumei Benth
Collards—Brassica oleracea var. acephala DC.
Coltsfoot—Tussilago farfara L.*
Columbine—Aquilegia spp.
Coneflower—Rudbeckia spp.
Coralbells—Heuchera sanguinea Engelm.
Coreopsis, perennial—Coreopsis lanceolate L.
Coriander— Coriandrum sativum L.
Corn (dent, field, flint, popcorn, sweet, white, yellow)—Zea mays L.
Cosmos—Cosmos spp.
Cotton—Gossypium spp.
Cowpea— Vigna unguiculata (L.) Walp. subsp. unguiculata
Cranesbill—Geranium spp.
Cress:
Garden—Lepidium sativum L.
Water—Nasturtium officinale R. Br.
Crested dogtail—Cynosurus cristatus L.
Crotalaria—Crotalaria intermeda Kotschy, C. juncea L., C. lanceo-
tata E. Mey., C. spectabilis Roth, and C. strata DC.
Crownvetch—Coronilla varia L.
Cucumber—Cucumis sativus L.
Cyclamen—Cyclamen europeum L.
Cypressvine—Ipomoea quamoclit L.
Dahlia—Dahlia spp.
Daisy:.
African—Dimorphotheca, aurantiaca DC.
African lilac—Arctotis grandis Thunb.
English—Bellis perennis L.
Painted—Pyrethrum spp.
Shasta—Chrysanthemum leucanthemum L.
Swan river—Brachycome spp.
Dallisgrass—Paspalum dilatatum Poir.
Dames rocket, sweet rocket—Hesperis matronalis L.
Dandelion— Taraxacum officinate Weber
Delphinium, annual, perennial—Delphinium spp.
Dichondra—Dichondra repens Forst.
Dill—Anethum graveolens L.
Dock, curly—Rumex crispus L. *
Dodder—Cuscuta spp.*
Dropseed, sand—Sporo bolus cryptandrus (Torr.) A. Gray
Dusty-miller—Centaurea candidissima Lam.
Eggplant—Solanum melongena L.
Endive—Cichorium endivia L.
Evening primrose—Oenothera biennis L. and 0. lamarckiana (0.
grandifiora Ait.)*
Fennel—Foeniculum vulgare Mill.
Fescue:
Chewings—Festuca rubra subsp. commutata Gaud.
Creeping red—Festuca rubra L.
Hair—Festuca tenuifolia Sibth.
Meadow—Festuca pratensis Huds. (syn. F. elatior L.)
Sheep—Festuca ovina L.
Tall—Festuca arundinacea Schreb.
Firebush, Mexican—Kochia spp.
Flax:
Common—Linum usitatissimum L.
Flowering—Linum grandiflorum Desf.
Perennial—Linum perenne L.
Forget-me-not—Myosotis spp. *
Foxglove—Digitalis spp.
Foxtail—Setaria spp.*
Gaillardia—Gafflardia spp.
Geranium—Geranium spp.
Geum—Geum spp.
Gilia—Gilia spp.
Globe amaranth-- Gomphrena globosa L.
Gloxinia, common—Sinningia speciosa (Lodd.) Hiern

Godetia—Godetia amoena Lilija and G. grandiflora Lindl.
Goodia clover—Goodia latifolia Salisb.*
Goosegrass--Eleusine indica (L.) Gaertn.*
Gourds—Cucurbita spp. and Lagenaria app.
Grama:
Blue—Bouteloua gracilis (H.B.K.) Lag.
Side oats—Bouteloua curtipendula (Michx.) Torr.
Guar—Cyamopsis tetragonoloba (L.) Taub.
Guineagrass—Panicum maximum Jacq.*
Hardinggrass—Phataris tuberosa var. stenoptera (Hack.) Hitchc.
Hawksbeard—Crepis spp. *
Heliopsis—Heliopsis spp.
Heliotrope—Hetiotropium spp.
Hemp—Cannabis sativa L.
Hibiscus—Hibiscus spp.
Hollyhock—Althaea roses (L.) Cay.
Hung tau—Vigna radiata (L.) Wilczek*
Hyacinth-bean—Dolichos lablab L.
Hyssop—Hyssopus officinalis L.
Indiangrass, yellow—Sorghastrum nutans (L.) Nash
Indigo—Indigofera cytisoides L.*
Indigo, hairy—Indigofera hirsuta L.
Iris, Japanese—Iris kaempferi Sieb.
Japanese lawngrass—Zoysia japonica Steud.
Jerusalem or Maltese cross—Lychnis chalcedonica L.
Jimsonweed—Datura spp.*
Jobs-tears—Coix lacryma-jobi L.
Johnsongrass—Sorghum halepense (L.) Pers.
Jute—Corchorus capsularis L. and C. olitorius L.
Kale—Brassica oleracea var. acephala DC.
Kenaf—Hibiscus cannabinus L.
Kidneyvetch—Anthyllis vulneraria L.*
Kohlrabi (knolkol)—Brassica oteracea var. gongylodes L.
Kudzu—Pueraria lobata (Willd.) Ohwi (syn. P. thunbergiana (Sieb. and
Zucc.) Benth.)
Lambsquarters, pigweed—Chenopodium spp. *
Lantana—Lantana camara L.
Larkspur:
Annual—Delphinium ajacis L.
Hybrids—Delphinium hybridum Steph.
Leadtree—Leucaena leucocephala (Lam.) de Wit*
Leek—A Ilium porrum L.
Lentil—Lens culinaris Medic.
Lespedeza:
Kobe—Lespedeza striata (Thunb.) Hook. and Am.
Korean—Lespedeza stipulacea Maxim.
Sericea or Chinese—Lespedeza cuneata (Dumont) D. Don
Siberian—Lespedeza hedysaroides (Pallas) Ricker
Striate—Lespedeza striata (Thunb.) Hook. and Am.
Lettuce—Lactuca sativa L.
Lily—Littium spp.
Linaria—Linaria spp.
Lobelia—Lobelia erinus L. and L. cardinalis L.
Locustbean (carob)—Ceratonia siliqua L.
Lotus, Indian—Nelumbo nucifera Gaertn.
Lovegrass, weeping—Eragrostis curvula (Schrad.) Nees
Lunaria, honesty—Lunaria annua L.
Lupine:
Annual types—Lupinus spp.
Arctic—Lupinus arcticus S. Wats.
Blue—Lupinus angustifolius L.
Russell hybrids—Lupinus polyphyllus Lindl.
White—Lupinus &bus L.
Yellow—Lupinus luteus L.
Maize (corn)—Zea mays L.
Mallow—MaIva spp.
Manilagrass—Zoysia matrella (L.) Merr.
Marigold:
African—Tagetes erects L.
French—Tagetes patula L.
Marjoram—Origanum majorana L.
Marvel of Peru, four-o'clock—Mirabilis jalapa L.
Matricaria—Malricaria spp.
Meadow foxtail—Alopecurus pratensis L.
Medic—Medicago spp.
Medic, black—Medicago lupulina L.
Mignonette—Reseda odorata L.
Milkvetch--Astragalus massiliensis Lam. (A. tragacantha) and A.
utriger Pallas*
Millet:
Foxtail (common, German, golden, Hungarian, Siberian)—Setaria
italica (L.) Beauv.
Japanese—Echinochloa crusgalli var. frumentacea (Link) W. F.
Wight
Pearl—Pennisetum americanum (L.) Leeke
Proso—Panicum miliaceum L.
Mimosa—Mimosa glomerata Forsk. *
Morningglory—Ipomoea spp.
Mullein, moth— Verbascum blattaria L. *
Muskmelon (cantaloup)—Cucumis melo L.
Mustard:
Black—Brassica nigra (L.) Koch*
India—Brassica juncea (L.) Czern.
Myosotis—Myosotis spp.
Nasturtium— Tropaeolum majus L.
Needlegrass—Stipa viridula Trin.*
Nemesia—Nemesia spp.
Nicotiana—Nicotiana spp.
Nigella—Nigella damascena L.
Nutgrass, nutsedge—Cyperus rotundas L.*
Oatgrass, tall—Arrhenatherum elatius (L.) Beauv. ex J. Presl and K.
Presl
Oats—Avena sativa L.
Okra—Hibiscus esculentus L.
Onion—Allium cepa L.
Onion, Welch—Allium fistulosum L.
Orchardgrass—Dactylis glomerata L.
Pakchoi—Brassica chinensis L.
Panicgrass—Panicum spp.*
Panicgrass, blue—Panicum antidotate Retz
Pansy— Viola tricolor L.
Parsley—Petroselinum crispum (Mill.) Nym. ex A. W. Hill
Parsnip—Pastinaca saliva L.
Partridgepea—Cassia fasciculata Michx.*
Paspalum—Paspalum spp. *
Path rush—Juncus bufonius L.*
Pea:
Austrian winter or field—Pisum sativum var. arvense (L.) Poir.
Garden—Pisum sativum L.
Rose or crown—Pisum umbellatum (P. sativum var. umbellatum
Ser.)
Peanut, Florida runner—A rachis hypogaea L.
Pechay (mustard)—Brassica juncea (L.) Czern.
Pennisetum—Pennisetum spp. *
Pennycress, fanweed—Thlaspi spp.*
Penstemon—Penstemon spp.
Pepper—Capsicum annuum L.
Pepper, red—Capsicum frutescens L.
Pe-tsai (Chinese cabbage)—Brassica pekinensis (Lour.) Rupr.
Petunia—Petunia hybrida Vilm.
Phacelia—Phacelia spp.
Phlox—Phlox drummondii Hook. and Phlox spp.

Physalis—Physalis spp.
Pimpernel—Anagallis spp.*
Pine—Pines spp.
Pinks, China—Dianthus spp.
Plantain—Plantago spp.*
Poppy:
California—Eschscholtzia californica Cham.
Corn, shirley—Papaver rhoeas L.
Iceland—Papaver nudicaule L.
Mexican tulip—Hunnemannia fumariifolia Sweet
Oriental—Papaver orientate L.
Portulaca—Portulaca grandiflora Hook.
Potato—Solanum tuberosum L.
Primrose--Primula spp.
Primrose, Chinese—Primula sinensis Lindl.*
Pumpkin—Cucurbita pepo L. and Cucurbita spp.
Pyrethrum—Chrysanthemum spp.
Radish—Raphanus sativus L.
Radish, Japanese—Raphanus sativus L.
Rape, annual and winter—Brassica napes L.
Redtop—Agrostis gigantea Roth
Rescuegrass—Bromus unioloides Kunth
Rhodesgrass—Chloris gayana Kunth
Rhubarb—Rheum rhaponticum L.
Rice—Oryza sativa L.
Ricegrass, Indian—Oryzopsis hymenoides (Roem. and Schult.) Ricker
Rose campion—Lychnis spp.
Rosemary—Rosmarinus officinalis L.
Roughpea—Lathyrus hirsutus L.
Rush—Juncus spp.*
Rutabaga—Brassica napes var. napobrassica (L.) Reichb.
Rye—Secale cereale L.
Ryegrass:
Annual— Lolium multiflorum Lam.
Italian—Lolium multiflorum Lam.
Perennial—Lolium perenne L.
Safflower—Carthamus tinctorius L.
Sage:
Mealycup—Salva farinacea Benth.
Sage—Salvia officinalis L.
Scarlet—Salvia splendens Sello ex Nees
Sainfoin—Onobrychis viciifolia Scop.
St.-Johnswort—Hypericum humifusum L. *
Salpiglossis—Satpiglossis sinuata Ruiz and Pay.

Salsify—Tragopogon porrifolius L.
Saponaria—Saponaria ocymoides L. and S. vaccaria L.
Savory—Satureja hortensis L.
Scabiosa—Scabiosa atropurpurea L. and S. caucasica Bieb.
Scotch-broom—Cytisus candicans Lam. (C. monspessulanus) and C.
scoparius (L.) Link*
Senna—Cassia bicapsularis L. and' C. ultijuga Rich.*
Sensitive plant—Mimosa spp.
Sesame—Sesamum indicum L.
Sesbania—Sesbania spp.
Silktree, mimosa—Albizia julibrissin Durazz.*
Sitao-- Vigna unguiculata subsp. sesquipedalis (L.) Verde.
Smartweed—Polygonum spp.*
Smartweed, water—Polygonum hydropiper L.*
Smilo—Oryzopsis miliacea (L.) Aschers. and Schweinf.
Snapdragon—Antirrhinum spp.
Sneezeweed, helenium—Helenium spp.
Snow-on-the-mountain—Euphorbia marginata Pursh
Solanum—Sotanum spp.
Sorghum, grain and sweet—Sorghum bicolor (L.) Moench
Soybean—Glycine max (L.) Merr.
Spinach:
Common—Spinacia oleracea L.
New Zealand—Tetragonia tetragonioides (Pall.) Ktze.
Squash:
Butternut—Cucurbita pepo L.
Winter—Cucurbita moschata Duchesne ex Poir. and C. maxima
Duchesne
Statice—Statice spp.
Stocks—Matthiola spp.
Strawberry—Fragaria x ananassa Duchesne
Strawflower—Hetichrysum monstrosum Hort.
Sudangrass—Sorghum sudanense (Piper) Stapf
Sun flower—Helianthus annuus L.
Swede—Brassica napus var. napobrassica (L.) Reichb.
Sweetciover:
White—Melilotus alba Desr.
Yellow—Melitotus officinalis (L.) Pall.
Sweetpea:
Annual—Lathyrus odoratus L.
Perennial— Lathyrus latifolius L.
Sweet-william—Thanthus barbatus L.
Switchgrass—Panicum virgatum L.
Thyme—Thymus vulgaris L.
Timothy—Phleum pratense L.
Tobacco—Nicotiana tabacum L.
Tomato—Lycopersicon esculentum Mill.
Trefoil:
Big—Lotus utiginosuo Schkuhr
Birdsfoot—Lotus corniculatue L.
Turnip—Brassica rapa L.
Vaseygrass—Paspatum urvillei Steud.
Velvetbean—Mucuna deeringiana (Bort) Merr.
Verbena— Verbena spp.
Vetch:
Common— Vicia sativa L.
Hairy— Vicia villosa Roth
Hungarian— Vicia pannonica Crantz
Monantha— Vicia articulata Hornem.
Narrowleaf—Vicia angustifolia (L.) Reich.
Purple— Vicia benghalensis L.
Woollypod—Vicia dasycarpa Ten.
Vinca, periwinkle— Vinca rosea L. and V. minor L.
Viola— Viola cornuta L.
Wallflower—Cheiranthus allioni Hort. and C. cheiri L.
Walnut—Juglans australis Griseb.
Watermelon—Citrullus lanatus (Thunb.) Matsum. and Nakai
Wheat:
Common— Triticum aestivun L.
Durum—Triticum durum Deaf.
Hard red spring—Triticum aestivum L.
Hard red winter—Triticum aestivum L.
Poulard—Triticum turgidum L.*
Red Winter Speltz—Triticum spelta L.
Soft— Triticum spp.
Soft red winter—Triticum aestivum L.
Spring—Triticum aestivum L.
White—Triticum spp.
Wheatgrass:
Fairway crested—A gropyron cristatum (L.) Gaertn.
Intermediate—Agropyron intermedium (Host) Beauv.
Pubescent—Agropyron trichophorum (Link) Richt.
Wheatgrass—Continued
Slender—Agropyron trachycaulum (Link) Malte ex H. F. Lewis
Standard crested—Agropyron desertorum (Fisch. ex Link) Schuh.
Tall—A gropyron elongatum (Host) Beauv.
Western—Agropyron smithii Rydb.
Wild rye:
Blue—Elymus glaucus Buckl.*
Canada—Etymus canadensis L.
Russian—Elymus junceus Fisch.
Willow, weeping—Salix babylonica L.
Zinnia—Zinnia spp.
Zoysia (see Japanese lawngrass and manilagrass)
Bromus polyanthus Scribn. *

Dioclea pauciflora Rusby*
Hovea linearis (J. E. Smith) R. Br.*
Kennedya apetala Loddiger*
Medicago orbicularis (L.) Bartal.*
Melilotus gracilis DC.*
Insects
Beetle:
Flat grain—Cryptolestes pusillus (Schönherr)
Khapra—Trogoderma granarium Everts
Sawtoothed grain—Oryzaephilus surinamensis (L.)
Borer, lesser grain—Rhyzopertha dominica (F.)
Cadelle—Tenebroides mauritanicus (L.)
Chalcid—Bruchophagus spp.
Grain moth, Angoumois—Sitotroga cerealella (Oliver)
Weevil:
Granary—Sitophilus granarius (L.)
Rice—Sitophilus oryzae (L.)
Animals
Lemming, collard—Dicrostonyx spp.
Mouse—Muridae spp.
Rat—Rattus spp.
Squirrel—Sciurus spp.
CONVERSION TABLES FOR TEMPERATURES AND

MEASURES
Temperature Conversion 12
The numbers in the center of each column refer to the temperature in
centigrade or Fahrenheit. They can be converted to either scale. If
converting Fahrenheit to centigrade, the equivalent temperature is on
the left in each column. If converting centigrade to Fahrenheit, the
equivalent temperature is on the right.
Measures of Weight
Measures of Length
Measures of Area
Measures of Volume
Measures of Capacity
LITERATURE CITED
ANONYMOUS.
1942a. SEED STORAGE PROBLEMS. Puerto Rico Agr. Expt. Sta. Ann. Rpt.
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1942b. DURATION OF VIABILITY OF SEEDS. Gard. Chron. 111: 234.
1959. SEEDS BAGGED TO KEEP. Agr. Res. 8: 10-11.
1968. 550-YEAR OLD SEED SPROUTS. Sci. News 94: 367.
ABDALLA, F. H., and ROBERTS, E. H.
1969a. THE EFFECTS OF TEMPERATURE AND MOISTURE ON THE INDUCTION OF
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and ROBERTS, E. H.
1969b. THE EFFECT OF SEED STORAGE CONDITIONS ON THE GROWTH AND YIELD OF
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ABDUL-BAKI, A. A., and ANDERSON, J. D.
1972. PHYSIOLOGICAL AND BIOCHEMICAL DETERIORATION OF SEEDS. In Koz-
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AGENA, M. U.
1961. UNTERSUCHUNGEN UBER KALTEEINWIRKUNGEN AUF LAGERNDE GETRIE-
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ALTSCHUL, A. M., KARON, M. L., KAYME, L., and HALL, C. M.
1946. EFFECT OF INHIBITORS ON THE RESPIRATION AND STORAGE OF COTTON
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AMERICAN SOCIETY OF AGRICULTURAL ENGINEERS.

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ANDERSON, J. A., and ALCOCK , A. W., eds.
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1946. THE INTERNAL INFECTION OF COTTON SEED AND THE LOSS OF VIABILITY IN
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ARNOLD, J. R., and LIBBY, W. F.
1951. RADIOCARBON DATES. Science 113: 111-120.
ASSOCIATED SEED GROWERS, INC. (ASGROW SEED Co.).
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1954. RULES FOR TESTING SEEDS. Assoc. Off. Seed Anal. Proc. 44: 31-78.
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ATKIN, J. D.
1958. RELATIVE SUSCEPTIBILITY OF SNAP BEAN VARIETIES 'TO MECHANICAL IN-
JURY OF SEED. Amer. Soc. Hort. Sci. Proc. 72: 370-373.
ATKINS, W. R. G.
1909. THE ABSORPTION OF WATER BY SEEDS. Roy. Dublin Soc. Sci. Proc. 12:
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1957. DIE SAMEN LANDWIRTSHAFTLICHER KULTURPFLANZEN IM GRUNDSTEIN
DES CHAMALIGEN NURNBERGER STADTTHEATERS UND IHRE KEIMFAHIG-
KEIT. Ztschr. f. Acker- u. Pflanzenbau. 103: 454-472.
AUSTIN, R. B.
1972. EFFECTS OF ENVIRONMENT BEFORE HARVESTING ON VIABILITY. In Rob-
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AVERY, A. G., and BLAKESLEE, A. F.
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BABBITT, J. D.
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BACCHI, 0.
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BAILEY, C. H.
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274.
BAINER, R., and BORTHWICK, H. A.
1934. THRESHER AND OTHER MECHANICAL INJURY TO SEED BEANS OF THE LIMA
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BAKER, D., NEUSTADT, M. H., and ZELENY, L.
1957. APPLICATION OF THE FAT ACIDITY TEST AS AN INDEX OF GRAIN DE-
TERIORATION. Cereal Chem. 34: 226-233.
BAKER, K. F.
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1963. IMPORTANT EQUIPMENT FOR DRYING SEEDS IN NORTH AMERICA. Inter-

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BARROW, E. H. R.
1915. THE HEATING OF COTTONSEED---ITS CAUSES AND PREVENTION. Indus.
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BARTON, L. V.
1932. EFFECT OF STORAGE ON THE VITALITY OF DELPHINIUM SEEDS. Boyce
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1939. STORAGE OF SOME FLOWER SEEDS. Boyce Thompson Inst. Contrib. 10:
399-427.
1941. RELATION OF CERTAIN AIR TEMPERATURES AND HUMIDITIES TO VIABILITY

OF SEEDS. Boyce Thompson Inst. Contrib. 12: 85-102.
1947. EFFECT OF DIFFERENT STORAGE CONDITIONS ON THE GERMINATION OF

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1953. SEED STORAGE AND VIABILITY. Boyce Thompson Inst. Contrib. 17: 87-103.
1954. EFFECT OF SUBFREEZING TEMPERATURES ON VIABILITY OF CONIFER SEEDS

IN STORAGE. Boyce Thompson Inst. Contrib. 18: 21-24.
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1960b. LIFE SPAN OF FROST-DAMAGED CORN SEEDS. Boyce Thompson Inst.

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1966a. THE EFFECT OF STORAGE CONDITIONS ON THE VIABILITY OF BEAN

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1966b. VIABILITY OF PYRETHRUM SEEDS. Boyce Thompson Inst. Contrib. 23:

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BAUTISTA, G. M., and LINKO, P.
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BEAL, W. 3.
1885. THE VITALITY OF SEEDS BURIED IN THE SOIL. Soc. Prom. Agr. Sci. Prof. 6:
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INDEX
AGRICULTURE HANDBOOK 506, U.S. DEPT OF AGRICULTURE

Principios y Practicas de Almacenamiento de Semillas

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PRINCIPLES AND PRACTICES

By Oren L. Justice and Louis N. Bass

Agriculture Handbook No. 506

On January 24, 1978, four USDA agencies—Agricultural Research

This publication was prepared by the Science and Education Admin-

Washington, D.C. Issued April 1978

apply equally to all seed storage. Longtime storage of genetic stocks is

The Seed, a Fragile, Living Organism

jected to the numerous mechanical and handling processes from harvest

From Harvest to Sowing

Decline and Death of Seeds

different times. For example, certain enzymes remain functional for a

SEED FACTORS THAT AFFECT STORAGE LIFE

live. Different percentages have been used for different purposes

Tonkin (1967) cheeked Brett's work by using large numbers of samples

MacKay and Tonkin (1S67) showed positive correlations between

Seed Structure and Composition

on the growth of mold, suggesting that the increased lifespan of cereal

likely to be extensively damaged. Size, arrangement of essential seed

C at moisture contents of 7-9 percent. Evaluation tests made alter 4

Panicum maximum var. Trechoglume seed increased in germination

hours lost 20-percent germination, but seed with 44-percent moisture

Adams, 1969). These studies showed that threshing or combining

Toole (1950) attempted to compare the germinability of slightly

ends at a survival level of 90-75 percent, and deterioration then

EFFECTS OF STORAGE ENVIRONMENT ON SEED

perature and moisture content to the period of seed viability of certain

TABLE 1. —Effect of various temperatures on germination of several

TABLE 2. —Effect of different temperatures on germination of

TABLE 3.—Effect of various temperatures on duration of seed germi-

Temperatures Below Freezing

Apparently seeds of barley, rye, and wheat are nearly as sensitiva as

Imbibed decorticated seeds had a considerably lower moisture content

Seed Moisture Content and Relative Humidity

consequently moisture content of the seeds on their storage life. They

(1957) found adsorption equilibrium values for corn and wheat to be

Relationship Between Relative Humidity and Seed Moisture

TABLE 5.—Equilibrium moisture content of vegetable seeds at various

TABLE 6.—Adsorbed equilibrium moisture content of grain seeds at

The sigmoid chape of a typical curve relating relative humidity to

TABLE 7 .—Equilibrium moisture content of seeds of grasses and

TABLE 8.—Equilibrium moisture content of seeds of grasses and

this disparity except possibly "the humidity readings in these chambers

increase in temperature--wheat 0.65 (Gane, 1941), 0.43 (Gay, 1946),

relative humidity is deereased below about 54 percent, the seeds at 49°

Rate of Moisture Absorption and Movement

moisture absorption of two cultivara of wheat at relative humidities

jute bags. The seeds had been stored at five temperature-relative

relative humidity of the storage atmosphere was decreased below 40

nation capacity had dropped by 5 percent and vigor was seriously

Interrelationship of Temperature, Seed Moisture

TABLE 11.—Germination of 2 lots of hempseed stored in sealed con-

terioration of seeds stored at ambient temperature and the same

Vacuum and Gas Storage

See footnotes at end of table.

TABLE 14.—Germination of aster seeds after storage at 12 tempera-

remained unchanged through 31 months, those at 50° in nitrogen

Confirmation Studies at the National Seed Storage Laboratory

Safflower seeds must be dried to 4-percent moisture (table 15) for

No atmosphere was consistently better than all others for preserving