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Plant, Cell atid Environtnent (1978) 1, 101-119.

The physiology of seed hydration and dehydration, and the

relation bet\A/een water stress and the control of germina-
tion: a review
T. W. HEGARTY Scottish Horticultural Research Institute, Invergowrie, Dundee

Received 8 January 1978; accepted for publication 30 January 1978

Abstract. Current interest in seed hydration treattnents tion process tliere is no doubt; the mechanisms of the
for the improvement of level, rate and uniformity of effect are, on the other hand, far from clear.
gennination or field emergence has revealed how little In addition, seed responses at different hydration
is Imown of the physiology of getmination control levels can vary widely, from germination on the one
under water stress. This review surveys briefly the re- hand to deterioration and death on the other, whilst
sponses of seeds held at different hydration levels, from some seeds can remain dormant and viable in the soil
normal germination in a free water supply to seed for many years while fully hydrated. It is the intention
activation without germination under slight moisture in this review to consider many of these responses to-
stress, seed deterioration at greater moisture stress and gether to see whether it is possible to build a more
the damage that can be caused to seeds in very dry con- comprehensive picture of the relation between seed
ditions, as well as the responses to subsequent dehydra- physiology and the degree of seed hydration. Of particu-
tion. Inhibition of germination, though not of seed lar interest are the precise responses of seeds to differing
activation, at certain levels of water stress is likened to levels of hydration; the effects of subsequent dehydra-
various forms of dormancy, and the mechanism govern- tion ; analogous or related situations that may yield rele-
ing the initiation of cell elongation is suggested as the vant infortnation on control mechanisms of seed
possible key to control over germination. Several lines germination under moisture stress; and whether non-
of evidence on cell membrane integrity and action, and germination under moisture stress is an active or passive
their responses to external factors, point to tlie role that response to the external environment.
the membrane may play in cell elongation (and hence This is not intended to be a comprehensive literature
germination) eontrol, and membrane integrity may review. Rather, I have used, where possible, references to
also be associated with the transition between seed general treatises or reviews which themselves include
deterioration at one hydration level and seed activation many of the references to original work; references to
and repair at slightly higher hydration levels. Seed activa- specific pieces of work are used where no review exists
tion witliout germination is also considered in an eco- or where specific poitits are being tnade. Consideration
logical context. is limited to those species in which the organ of protru-
sion from the seed is the radicle rather than tlie coty-
ledons or hypocotyl.
(1) Introduction
(2) Historical perspective of hydration treatments
There has recently been a considerable upsurge of in-
terest in the use of pre-sowing seed treatments involving Much of the work of the last century was reviewed by
the full or partial hydration of seeds, sometimes fol- Kidd & West (1919). They concluded from the results
lowed by subsequent dehydration, which result in an published that seeds swollen in water and sown still in
increase in the rate, uniformity or level of seed germina- the moist condition germinated more rapidly than un-
tion.* These seed treatments for improved performance treated seeds; tliat seeds soaked in a 'minimum' amount
have been described, amongst many other kinds of seed of water and afterwards dried slowly at ordinaty tem-
treatment, by Heydecker & Coolbear (1977). That the peratures imbibed water and developed more rapidly
treatments have substantial effects on the seed germina- when again allowed to take up water and germinate,
than untreated seeds; but that seeds dried rapidly after
* In this paper 'seed' is used to describe all types of propagule, initial soaking germinated tnore slowly than untreated
'germination' is used to describe the emergence of the radiele seeds. Particular reference was made to the conditions
from the seed, and the 'germination process' is that series of
events culminating in germination itself. of seed soaking such as the quantity of water, duration
and temperature of the treatment, and to the apparent
Correspondence: Or T. W. Hegariy, Scottish Horticultural
Researeh Institute, Invergowrie, Dundee. lack of effect of salts added to the soaking solutions.
Their own experiments (Kidd & West, 1918) indicated a
1978 Blaekwell Seientific Publications beneficial response to soaking in some cases but deleteri-


ous effects in others. Chippindale (1934) investigated their apparent advantages the method of seed priming
the effects of seed soaking on grasses, and mentioned that is now being used for commercial development in
that the practice was formerly carried out frequently by Britain (based on the work of Darby & Salter, 1976) has
farmers, but not in 'modern' agriculture. He concluded been adapted to use salt solutions in place of the mote
that the effects found could not be generalized over viscous polyethylene glycol solutions.
species, even when closely related. This adaptation is, in fact, very similar to the method
A move towards making a more defined use of seed used in the U.S.A. (possibly based on the work of Ells,
hydration treatments was made by Le\att & Hamm 1963) and described briefly by Harrington (1969) which
(1943) who were concerned to ftnd a way to 'mature' involved soaking seeds in aerated hypertonic solutions of
seeds of Taraxacum kok-saghyz Rodin, before sowing. salts which just, or almost, prevented radicle emergence
They speculated that it should be possible to increase from the seed. After the hypertonic soak, seeds were
the moisture content of the seeds sufficiently to pennit dried back before sowing, but Salter & Darby (1976)
the necessary physiological changes, but insufftciently to proposed their method as a means of giving synclironous
permit gertnitiation, and then dry back the seeds so that germination before sowing the seeds directly in the field
they could be sown in the usual way. Accordingly, they using a fluid carrier.
first tried keeping seeds in contact with dextrose solu-
tion, used as an osmoticum permitting water uptake to
a controlled level, before drying them back. Having
obtained beneficial results, they went on to try salt (3) Effects of imposed treatments
solution (e. -2.7 MPa) or 0.5 M KNO3 solution (c. -2.0 Conclusions regarding seed hydration treatment effects
All the methods hastened subsequent germitiation, the can differ little today from those reached by Crocker &
optimum treatment incorporating molar (M) dextrose Barton (1957) or, indeed, by Kidd & West (1919)
solution (c. -2.7 MPa) or M/2 KNOa solution (c. 2.0 except that it is now recognized that the effect of salts
MPa), with optimum duration dependent upon the tetn- in the soaking medium can be important. The effect
perature. There was no loss of benefit in dehydrated depends upon the degree of seed hydratioti, tempera-
seeds stored for 4 weeks. ture and duration of the treatment, the level of aera-
To a considerable extent this work seems to have re- tion, density of seed mass and whether (possible how)
mained utirecognized, and only 20 or so years later, the seeds are dried back. Treatments used involve soak-
possibly after the publication of a review of Russian ing seeds in water or on surfaces with a free or restricted
work on seed 'hardening' (May, Milthorpe & Milthorpe, supply of water, or allowitig them to absorb water from
1962) was there an increase in interest in the use of vapour. The treatment may also have involved subse-
pre-sowing treattnents using a restricted quantity of quent dehydration, possibly with several cycles of
water. The Russian work was concerned with inducing hydration and dehydration. The treatments and their
drouglit tolerance during subsequent plant growth, and effects have been reviewed by Heydecker (1974) and by
consisted essentially of variations on the use of one or Heydecker & Coolbear (1977). Different investigators
more cycles of seed hydration in a restricted quantity of have reported favourable, neutral and unfavourable
water followed by dehydration after a specified time. It results. Generally, however, germination or field etner-
was considered that as germination proceeded the resis- gence is more rapid after treatmetit, especially at low
tance of the embryo to dehydration decreased and the temperature (Bremer, 1928; Chippindale, 1934), the
likelihood of permanent injury from drying increased effect being more pronounced if the seed is not dried
whereas the degree of 'hardening' induced was likely to back (Heydecker, 1974). Germination can also be more
be greater the more advanced the embryo at the time of uniform or synchronous (Heydecker, 1974; Salter &
dehydration. Datby, 1976). Germination or field emergenee levels
Meydecker (1974), in a paper reflecting to some may also be improved. The increases in field emetgence
extent the philosophy expressed 30 years earher by of carrot {Daucus carota E.) described by Bremer (1928)
Levitt & Hatnm (1943), questioned whether there was and Currali & Salter (1973) may have resulted sitnply
any physiological advantage in drying back seeds after from the increased rapidity of establishment of treated
treattnetit, or whether this was done simply for con- seeds in the field, but the results of Chippindale (1934)
venience. He also recognised that the salt solutions for grasses, Eevitt & Hamm (1943) for Taraxacum kok-
used in seed soak treattnents to 'feed' seeds may simply saghyz, Hegarty (1970) and Heydecker, Higgins &
have been having an osmotic effect, and he described Turner (1975) for carrot, Sachs (1977) for watermelon
the development of the method of seed 'pritning' using {Citrullus lutwtus (Thunb.) Matsum. & Nakal.), and
polyethylene glycol solutions, with solute potentials Khan & Knypl (1977) for lettuce (Lactuca sativa L.),
generally between -1.0 and -1.5 MPa, which bring celery {Apium graveolens L.), onion (AUium eepa L.)
seeds to the 'brink' of germination apparently by allow- and soybean (Glyeine max Merr.) showed clearly that
ing etnbryo development but not germination. Poly- percentage germination at low temperature ean be im-
ethylene glycols were selected because they appeared to proved, whilst Gelmond (1965) for leek (AUium porrum
be true osmotica, free from the complications of ionic or E.) and Heydeeker et al. (1975) for lettuce demon-
molecular uptake (as in the case of sugars) or of specific strated that germination at high temperature can also be
ion effeets (Uhvits, 1946; Redmann, 1974), but despite inereased. The requirement for alternating temperature

in Dactylis glomerata E. can also be removed by a pre- the cotyledons in cotton possibly acting as reservoirs of
hydration treatment (Haight & Grabe, 1972). In addi- water for the embryo. The physiology of seeds at hydra-
tion to overcoming tetnperature restrictions, seed tion levels leading to normal germination is considered
soaking treatments have been shown to affect subse- in Section 4.1.
quent germination under osmotic stress or in saline Seeds may be hydrated via either the liquid or vapour
conditions. Hajison (1973) repotted faster germination phases and both have been used to bring seeds to equi-
of pretreated wheat (Tritieum aestivum E.) seeds in a librium water contents below those that pennit germina-
solution of 0.3 MPa solute potential compared to tion. (Equilibration at very high relative hutnidities
untreated seeds, and both rate and level of gennination (RH)* is considered separately in Section 4.6). In seed
of wheat in sahne eonditions have been iticreased by priming (Heydecker, 1974), an osmotic solution is used
seed treatments (Chaudhuri & Wiebe, 1968; Idris & to control water uptake whereas in seed 'hardening' or
Aslam, 1975). Gray & Steekel (1976) found that a pre- 'advancing' (Austin, Eongden & Hutcliinson, 1969), a
soak in phosphate buffer solution improved both the known quantity of water is added to the dry seeds to
percentage germitiation of lettuce in 0.16 M NH4NO3 bring them to the required water content. In both these
solution (c. -0.7 MPa) and seedhng emergence in soil treatments the seeds are sufficiently liydrated to permit
to which NH4NO3 had been added. Pre-soaking wheat seed activation without deterioration, but if seeds are
seeds in water immediately before planting has also hydrated to ratlier lower moisture contents ('damp'
been found to increase seedling emergence from salt- seeds) they deteriorate rapidly. Seed storage deteriora-
containing soil (Eyles & Fanning, 1964) and seed germi- tion work often uses vapour equilibration to eontrol the
nation on saline media (Chaudhuri & Wiebe, 1968). level of seed hydration. The physiology of activated
Villiers & Edgcumbe (1975) have shown that inter- seeds is considered in Section 4.4. and that of 'damp'
mittent periods of imbibition followed by dehydration seeds in Section 4.5.
can prevent deterioration of stored lettuce seed, and The extent of seed hydration at a given water poten-
Basu et al. (1975) have reported (without presenting tial is dependent upon the character of the particular
data) that a hydration-dehydration treatment extended seeds, varying with the species and seed constituents
the subsequent storage life of a wide range of seeds (Owen, 1956; Crocker & Barton, 1957; Roberts &
under ambient conditions or during accelerated ageing Roberts, 1972) as well as with individuals within a seed
under different tetnperature and humidity regitnes. stock (see Section 5). Harrington (1973) has described
a generalized and traditional view of the equilibrium
water relations of seeds. If the seed moisture content is
(4) Seed physiology in relation to hydration level
liigh enough (over 43%t), non-dormant seeds will germi-
Dry seeds have water potentials of less than 100 MPa nate; in the range from 22 to 43%, heating due to micro-
and, except in the case of 'hard' seeds, rapidly imbibe organisms will occur if oxygen is present, resulting in
when placed in good contact with iree water, though the rapid death of the seeds; in the range from about 11%
amount of water taken up depends upon the particular (oily seeds) or 15% (starchy seeds) to about 22%,
seed constituents, upon the individual seed and upon storage fungi grow actively and destroy the seed embtyo.
the tetnperature. This first, rapid phase of imbibition is These last values represent moisture contents in equi-
apparently a purely physical process, occurring equally librium at 20C to 25C with RH's of 65-70%) at the
in live or dead seeds (Mayer & Poljakoff-Mayber, 1975). drier end, and 85-90% at the wetter end of the range
Excess of water may prevent tioimal getmination (see (Christetisen, 1973). Thus this view essentially proposes
Section 4.2, below) but if there is adequate aeration the three effective ranges: fully itnbibed seeds which will
seeds will germinate. However, full germination does germinate; damp seeds that deteriorate in association
not require full seed hydration but can occur over a with microorganisms; and dry seeds. This simple view
range of water potentials and hence degrees of seed fails to represent the data adequately, and instead
hydration, the range being dependent on the particular Hegarty (1977b) has proposed five ranges of seed hydra-
species involved (Doneen & MacGillivray, 1943; Hunter tion which take into accoutit in addition seed activation
& Eriekson, 1952; McGitinies, 1960; Kauftnann, 1969; without gennination, and seed damage in very dry con-
Hegarty, 1976). In addition, a continuous supply of ditions, the latter being discussed in Section 4.3.
water to the seed is not essential for getmination to pro- As an addendutn to the introduction to tliis Section
ceed. MeDonough (1975) has shown for a selection of it is wotth etnphasizing the nature of the relationship
grasses, legumes and herbs that itnbibed seeds can be between RH, water potential and seed water content.
removed from their souree of water supply before germi- At 15C, a water potetitial of -0.15 MPa is equivalent
nation starts, but can still germinate, thus demonstrating
* The preferred units for describing the moisture eontetit of
elearly that water held within the seed structure can be air are g m"' rather than % RH. The former is, however, tem-
used to pertnit extension growth. Indeed, Stiles (1948) perature dependent, and as temperature data have not been
had shown earlier that different orgatis within the seed included in all the papers in which RH figures are given, conver-
hydrate to different extents, with the embryos in maize sion of % RH to g m '' is not possible in all cases. For consis-
(Zea mays E.) and cotton (Gossypium hirsutum L.) tency, % RH has been used throughout.
taking up water to a far greater extent than the other t Seed tnoisture eontents are expressed on a dry weight basis
and data from papers have been adjusted to this base where this
seed structures, and witli the endosperm in maize and has been necessary.

to 99.9% RH; -1.5 MPa to 98.9% RH; -15.0 MPa to through effects on reactivation and, in the case of phase
89.5% RH; and -150 MPa to 33% RH (Slatyer, 1967). C, on catabolism.
In very approximate terms a seed may contain half as The rapidity of reactivation of pre-existing systems
much water again at 0 MPa as at 1.5 MPa (Hegarty, has been emphasised by the work of Marcus, Feeley &
1977a) so that at equilibrium the seed would be two- Volcani (1966) who found in wheat embryo that the
thirds hydrated in an atmosphere of 99% RH, the rise in protein-synthesising capacity lagged only 10
remaining one-third of potential water uptake being minutes beliind water uptake at room temperature. The
associated with a change of only 1% RH (if hydration independence and interdependence of the relationsliips
by vapour and liquid can be equated). in phases A, B and C of the germination process have
been described by Fujisawa (1966) in an investigation of
the action of various drugs on the development and
(4.1) Seeds hydrated sufficiently to permit germination growth of decotyledonized radish {Raphanus sativus L.)
During the germination process, water uptake occurs embryos, and by Sutcliffe & Bryant (1977) in their
essentially in three phases, with a more or less distinct description of seed germination in the pea {Pisum
static or lag phase separating two phases of rapid water salivum L.). From these descriptions it is clear that
uptake, the first associated with initial imbibition and developments in phases B and C are dependent on pro-
the second with radicle growth. The extent of the lag cesses occurring in their preceding phases, and in radish
phase is dependent upon the species, individual seed and it appeared that even witliin a single phase (B), progress
temperature (Berrie & Drennan, 1971; Hegarty, 1977a), was dependent upon processes occurring in an early
but although water uptake may be negHgible during period of that phase. However, respiratory activity in
this phase, the seed is not inactive. phase B was not affected in conditions wliich inhibited
respiration in phase C.
According to Brown (1972), it used to be commonly
accepted that the stage up to the rupturing of the seed For continuing radicle growth there must be concur-
coat by the radicle was more or less passive, with events ring cell division and cell extension, although their onset
determined by the physical properties of the system. may not be simultaneous (Berlyn, 1972). Rogan &
Instead we should now recognize the pre-germination Simon (1975), using their own data and the results of
phase as including: an activation of enzymes that are other workers, describe an initial slow increase in root
present in the dry seed; a change in the structure of the length in broad bean {Vicia faba L.), barley {Hordeum
cytoplasm that sustains metabolic activity; a synthesis vulgare L.), pea and French bean {Phaseolus vulgaris
of enzymes both before and after active growth has L.) which is followed by a phase of rapid growth co-
begun; and a change in the metabolic pattern as a inciding approximately with the onset of mitosis.
result of a change in the quantitative relations between According to Sutcliffe & Bryant (1977), increase in the
enzymes in the growing system. volume of the radicle in pea causes it to emerge from the
This pre-germination phase has itself been subdivided testa before the start of the phase of rapid growth (tliis
into phases by a number of different authors. Evenari latter involving a large increase in DNA and RNA con-
(1961) describes four phases (imbibition; activation; tents of the axis accompanied by rapid cell extension).
mitosis; and protrusion) whilst Berlyn (1972) also sug- This coincidence of cell division and rapid cell elonga-
gests four phases, though with a slight difference in tion has also been found in oat {Avena sativa L.) and
emphasis (imbibition; hydration and activation; cell tomato {Lycopersicon esculentum Mill.) (Berrie &
division and extension; and protrusion of the embryo Drennan, 1971) and in lettuce at 26C (Evenari et al.,
from the seed). Perhaps more closely correlating with 1957). However, Haber & Luippold (1960a), whilst con-
the pattern of water uptake and respiration in seeds is firming the results for lettuce at 26C, were able to show
the less rigid description of the germination process by that cell division and cell elongation were not necessarily
Ching (1972; 1973) in terms of three distinct but over- synchronous, but could be delayed with respect to each
lapping phases. The first (A) involves the hydration and other by low temperature (10C delaying mitosis) or
reactivation of existing systems when there is likely to by osmotic stress (0.4-0.5 M mannitol, 1.0 to 1.3
be a start to basal metabolism, an increase in ATP con- MPa, delaying elongation). Under extreme circumstances
tent for various synthetic activities following imbibition, (heavily irradiated seeds) plantlets of some species
and the provision of substrate for respiration and protein capable of CO2 fixation and dry weight increase can
synthesis. The second phase (B) involves synthesis of grow, even thougli mitosis has been completely sup-
enzymes and organelles for catabolism of reserves and pressed (Berlyn, 1972). TIius radicle protrusion from the
occurs mostly in the storage organs, with the activity seed may be due to cell elongation in combination with
or quantity of the protein-synthesizing machinery, mito- cell division or to cell elongation alone wliich can result
chondria, glyoxysomes, enzymes, coenzymes, cofactors, in 'false' germination, a phenomenon that is considered
substrates and other factors increasing during and after in Section 8,below.
imbibition. The third phase (C) involves synthesis of Respiratory patterns during the germination process
new cellular components and is associated with radicle are complex and have been discussed by Roberts (1969,
emergence from the seed, using substrate transported 1973a) and, briefly, by Mayer & Shain (1974). In dry
from the storage organs. Both phases B and C are likely seeds mitochondria have been found to be poorly dif-
to be affected by environmental influences, possibly ferentiated and lacking in malate dehydrogenase, but as

imbibition proceeds they become rich in protein and ethanol are due to a deleterious 'fluidizing' of the cell
Upid, and biologically active (Nawa & Asahi, 1971). A membranes. Certainly, the damage associated with the
shift in respiratory pattern 4-8 h after imbibition has leaching of solutes at higli and low temperatures or in
been found in soybean, oat, lettuce, corn, tomato and the presence of excess oxygen, described above, would
pea, from predominantly alternate respiration to a indicate a general role of membrane involvement in
cyanide-sensitive respiration (Yentur & Leopold, 1976), soaking injury.
and Howell (1961) reported alterations in metabolic
characteristics of soybean cotyledon mitochondria
during the course of the germination process. Toole et (4.3) Very dry seeds
al. (1956) have suggested that in a physiological sense, Over-desiccation can damage seeds (Harrington, 1972;
the start of germination depends upon coupHng respira- Ching, 1973; Woodstock, Simkin & Schroeder, 1976).
tion to growth. However, whilst radicle growth may be Harrington (1972) suggests that water in seeds can be
linked with increased respiration, respiration can pro- described in terms of three categories: unbound, mobile,
ceed at a substantial rate in the absence of growth, i.e. free, capillary or solution water; multilayer, chemi-
in seeds that for one reason or another cannot germinate sorbed, hydrogen-bonded, intermediate water; and
(Ching & Foote, 1961; Chen & Varner, 1970; Hegarty, monolayer, localized, polar-site, bound water. As seeds
1977a) and long before growth begins, energy is coupled are dried, first the free, then the intermediate water is
to endergonic metabolic processes (Haber & Luippold, lost, and below 25% RH the monolayer water is progres-
1960b). However, the extent to which this respiration is sively lost from the macromolecules, removing a layer
associated with embryo development or repair mechan- that is protective against oxidative processes. The most
isms, and the extent to which respiration is uncoupled, serious oxidation is the autoxidation of lipids which can
'awaiting' growth, is not clear. continue as a chain reaction and which results in the
production of free radicals which are very reactive even
in seeds of low moisture content. Seeds contain natural
(4.2) Soaked seeds antioxidants but these are used up during storage as
Seeds immersed in water imbibe, may lose a wide range oxidation occurs and no more are produced in very dry
of solutes and, in time, may germinate or suffer damage. seeds.
Although many seeds will not germinate under water, Whilst it might be expected that withdrawal of mono-
some certainly can, in a very restricted supply of oxygen layer water would itself affect macromolecular structure
(Carr, 1961). In an extensive investigation, Morinaga and function, the free radicals produced can damage
(1926) placed seeds in Erlenmeyer flasks filled with cellular membranes and can react destructively with
distilled or boiled water. He found that germination of macromolecules, destroying protein, DNA, and cell
18 species was virtually the same under water as on filter reproductive capacity (Harrington, 1972; Harman &
paper, 25 species showed reduced germination, and 34 Mattick, 1976).
species no germination. The ability to germinate under
water was associated more with small seed size than with
large seeds but was not related to phylogeny or to kinds (4.4) Seeds sufficiently imbibed to be activated but
of reserve material in the seeds. An atmosphere of oxy- unable to germittate (incompletely imbibed seeds)
gen rather than air in the flasks gave liigher levels of Data on the pliysiology of incompletely imbibed, non-
germination. dormant seeds are sparse and scattered, and the picture
According to Carr, when seeds arc soaked in water of what is happening in such seeds must to a large extent
they go through a period of anaerobiosis which, if too be inferred from work such as that of Heydeeker et al.
protracted, can cause injury, resulting in inability to (1975) and Salter & Darby (1976) that seeds can be
germinate or in poor germination and ready decay. The brought to the 'brink' of germination if under the ap-
state of tlie seed and the environmental conditions seem propriate osmotic control. The very rapid germination
to be important. For instance, dormant seeds appear on transfer to water, and the improved uniformity of
not to be damaged excessively by storage in water; germination imply that the germination process in these
increased injury in some species at 10 or 30C compared seeds has proceeded via one route or another until a
to 20C is associated with greater losses of solutes from particular block is reached (i.e. phases A and B are
the seeds; and many kinds of seeds are injured by soak- nearly if not altogether completed, thougli perhaps at a
ing in oxygenated water but may be protected by the slower rate than in water), and that the seeds within a
presence of hydrogen peroxide or, more readily, by single population reach tliis block at different rates but
carbon dioxide. In Frencii bean. Barton & McNab germinate relatively uniformly once the block is released.
(1956) found that oxygenation caused excessive water The data presented by Hagarty (1977a) on the
uptake in the seeds, and after extended (24 h) soaking respiration of calabrese {Brassiea oleraeea var italiea
in the presence of oxygen there was also excessive Plenck.) and carrot seeds when held in solutions of dif-
leaching from the seeds. ferent solute potentials showed that in the lag phase of
Crawford (1977) has suggested that seeds resistant to the germination process (phase B) the rate of oxygen
soaking injury regulate glycolysis so that there is mini- consumption was dependent on the degree of seed hy-
mal production of ethanol, and that the toxic effects of dration, but even at hydration levels which just pre-

vented germinafion, oxygen uptake rates were relatively minimizing the deterioration resulting at the moisture
higli. Malnassy (1971) found that the addition of cyclo- level of 'damp' seeds and that caused by excessive desic-
heximide (an inhibitor of protein synfhesis) to the seed cation. Bass (1975) suggests equilibrium with 30% RH
soak medium (ati aerated solution of NaCl at c. 1.0 as being best for many seeds, whereas Harrington (1972)
MPa) partly or completely negated the beneficial effect suggests equilibrium wifh 20-25% RH as the opfimutn.
of the seed soak on subsequent germination of pepper Changes that occur during storage and are associated
{Capsicum annum L.), celery and totnato seeds. He also with deterioration lead to delayed gennination, reduced
found isoenzyme differences in treated and untreated seedling growth rates, decreased tolerance of adverse
seeds, and quoted the work of Koehier (1967) showing germination conditions atid loss of germinability (Abdul-
that levels of RNA increased 80%, and peroxidase levels Baki & Anderson, 1972; Delouche & Baskin, 1973).
increased eightfold in tomafo seeds during the treat- However, the underlying causes of these chatiges are tiot
ment. High nucleic acid levels in 'primed' tomato seeds clear, tiot so much because no cytological and metabolic
were also reported by Coolbear & Heydecker (1975). changes have been detected, but because so tnany have
Using 'pdmed' seeds of lettuce, Klian et al. (1978) been recorded (Roberts, 1973b). Roberts described
have shown that, compared to untreated seeds, there are sotne of the tnore consistent observed changes in seeds
iticreases in the rates of ^H-uridine iticorporation into which include: cytological damage, including datnage to
RNA, polyribosome incorporation and '''C-leucine the chromosomes and ribosotnes sytnptomatic of a
incorporation into profeins; iticreased activities of sotne general degradation of nucleic acids, damage to other
enzytnes thougli not of others; and qualitative and organelles symptomatic of general degradation of hpo-
quantitative changes in soluble proteins and isoenzymes protein tnetnbranes, atid datnage fo the cell tnetnbratie
of esterase, 3-phosphoglyceraldehyde dehydrogenase, itself; alterations in the activity of enzytnes, particularly
and phosphatases. In addition, a 2-week osmotic treat- those concerned with redox activity which decrease in
ment at ]5C led to a complete disappearance of ab- activity, and sotne hydrolytic enzymes which increase in
scisic acid. activity; various changes in cell consfituenfs includitig
Subsequent reliydration after dehydration has re- profeins, fatty acids, reducing and tion-reducing sugars
sulted in higher respiration rates in treated compared to and growth inhibitors; and a decreased ability to incor-
untreated seeds (Koehler, 1967; Woodstock, 1969; porate isofopically labelled sugars and amino acids.
Malnassy, 1971), suggesting that substrates may be Some of these changes ate accelerated by micro-
accutnulated, or systetn activity may be enhanced in organisms, but Roberts considers that they can also
seeds that are just prevented from gennination. occur in their absence and in any case are compatible
In addition, cell division has been shown to occur in with the general increase in hydrolyfic activity.
carrot embryos at seed hydration levels that almost Accelerated ageing tests set out specifically to ac-
certainly would have prevented gennination (Austin ct celerate the processes of nonnal ageing by exposing the
al., 1969) and in embryos of lettuce seeds held in seeds to high RH and temperature for a specified period
mannitol solution (c. 1.3 MPa) which virtually pre- of titne. In general, the tnost usual conditions are 100%
vented germinafion (Haber & Luippold, 1960a), wliilst RH at 40-45C with exposure titnes of from 2-8 days,
cells in a metabolically active state were detected in but in some cases a less severe regime of 30C and 75%
radicles of Nemnphila mcnziesii Aggr. held in mannito] RH for periods of 6-24 weeks is used (Delouche &
solution of c. -2.0 MPa (Cruden, 1974). Baskin, 1973). Loss of vigour precedes loss of viability,
Thus considerable activity is possible in partially but fhe test itself is concerned only with nortnal gertni-
hydrated seeds, including, apparently, many or all of the nation after ageing, with high vigour seed lots showing
processes of phases A and B of fhe germination process, less reduction iti germination than low vigour lots.
and possibly even the operation of systems permitting There is much evidence that deferiorafion in 'datnp'
cell division, but not cell elongation co-ordinafed with seeds is a physiological process, but Christensen (1973)
cell division. It has not yet been shown, however, has argued that there is sufficient evidence fo itnplicate
whether the physiology of incompletely imbibed seeds fungal activity as a primary cause of seed deterioration,
is essentially similar to that of fully imbibed seeds up fo and he is careful to distinguish between the effects of
a particular point and then diverges, or whether the storage fungi, active in RHs of from 70-90%, from those
patterns of development in these fwo situations are quite of field fungi, active in excess of 95% RH. However,
distinct. Harrison & Perry (1976) have presented convincing
evidence for fhe role of physiological deterioration of
barley held in damp store. Deferioration increased with
(4.5) 'Dry'and 'damp'seeds iticreasing seed moisture content, and at most moisture
At room temperature 'dry' seeds of tnany species can be contents the start of decHne in vigour and viability
stored satisfactorily, but even af the optimum moisture clearly preceded any substantial invasion by fungi.
content for storage, seeds will deteriorate wifh age, the Deterioratioti in this case was measured in terms of
rate of deterioration (Roberts, 1972) and metabolistn reduced membrane integrity, atid both reduced respira-
(Bailey, 1940; Edwards, 1976) increasing as fhe seed tion and ability to synthesise a-atnylase during sub-
moisture content is raised. Thus the optimutn moisture sequent germination.
contenf for dry storage represents a balance between Loss of membrane integrity seems to be intimately

involved in the deterioration process (Koostra, 1973) out that absolute uniformity in moisture content is
and is probably a primary step in the sequence of events almost itnpossible to attain even in a stnall satnple of
during detetioratioti (Delouche & Baskin, 1973). seeds. Third, hydi'ation treattnents seetn to be particu-
larly setisitive to the content of the gaseous environ-
ment (Crocker & Barton, 1957; Heydecker, 1974) and
(4.6) Seed responses at very high relative humidities differences in oxygen or carbon dioxide concentrations,
If water potential predominantly controls water uptake especially around seeds held in confined atmospheres,
in seeds, one wotild expect sitnilar results of seed treat- could weD have influenced the results. Fitially, the
ment whether carried out in contact with a water- physiological response of the seed tnay be determined
supplying substrate or in controlled atmospheres. The by the rate of increase of hydration and by the absolute
exception to this would be if physical hydration were level of hydration when in equilibrium with saturated
different for some reason in the two situations, as has water vapour. If the rate of increase of hydratioti is slow,
been suggested by Shull (1916), or if the rate of supply the seed may go througli a protracted peiiod at hydra-
of water were importatit. Much of the work on seed tion levels that favour deterioration rather than activa-
'pritning' has involved the use of solutions in the range tion and repair, and although a higher tetnperature
1.0 to -2.0 MPa and, as has been pointed out earlier, would favour more rapid hydration, it would dso favour
an osmotic solution of -1.5 MPa at 15C is in equi- tnore rapid deterioration. In addition, seeds if truly
libriutn with a RH of c. 99%. Results of investigations equilibrated with saturated vapour may well be sub-
concerning the effects of saturated or nearly saturated stantially less hydrated than if itnbibed in contact witli
atmospheres on seeds are widely contradictory, with water in the liquid phase, especially if there are parts of
reports of seed germinatioti in some cases, activation the true seed or outer coverings that can retain quanti-
without germination in others, and deterioration to a tities of 'free' water which can be utilized by tlie seed
greater or lesser extent in a thitd group. for cell elongation during germination.
Equilibrium tiioisture contents attained at 100% Thus discrepancies reported on the effects of high
RH have been quoted for a range of crops (mainly RH on seeds could well relate to the absolute level and
cereals) by, for instance, Coleman & Fellows (1925) rate of hydration of the seeds, to the degree of tem-
and by James (1967) witli values ranging from approxi- perature control over the environtnent in which the
tnately 27% in flax {Einum usitatissimum L.) to 37% in seeds are held, and to the levels of carbon dioxide or
barley, the values quoted in the two papets agreeing oxygeti maintained in that environment. However, it
closely. Coleman & Fellows used a fiow of saturated air does seem possible that, in vapour, under carefully
rather thati a static systetn. No tnention was ttiade in controlled temperature conditions, seed activation and
either case of fungal growth or germitiation occurritig. germination can occur over a range of water potentials
Deterioration of seeds in saturated atmospheres has (Owen, 1952).
been described by Robertson, Lute & Gardner (1939),
Brewer & Butt (1950) and Griffin (1966) whereas other
reports describe germination (Black, 1959; Owen, 1952) (4.7) Conclusions
and improvemetits in subsequent seed gertnitiation Ouite distinct physiological processes occur at different
(Levitt & llatnm, 1943; Wilson, 1973; Heydecker, seed moisture contents, although they are not well
1974). Accelerated ageing treatment tnay involve high understood and are at best recognized through their
hutnidity storage and results in seed deterioration effects on seed constitution or on perfortnance during
(Delouche & Baskin, 1973). Fitially, Villiers (1974) germination. In 'very dry' seeds, datnage tnay be the
compared lettuce seeds hydrated in water vapour for 2 result of excessive removal of water frotn the seed; in
weeks befote sealed storage, with seeds kept fully im- 'dry' seeds, deterioration occurs but its rate is tninimal;
bibed but thermodormant and found that the former in 'datnp' seeds, both metabolism and deterioration ap-
showed deterioration whereas the latter did tiot. He parently increase with water content; at higher water
noted, however, that the vapour-equilibrated seeds had potentials there must be a transition point when repair
attained a moisture content of only 14%. reactions are activated or co-orditiated, or metabolism
Apart from the possible different responses of indi- becomes linked with seed activation, so that activatioti
vidual species or seed lots to being held in saturated and repair rather than deterioration are dominant.
water vapour, four explanations of these differences Substatitial etnbryo developtnent can occur in this state
seem possible. First, it is extremely difficult to main- of 'incomplete' imbibition. Only at higlier water poten-
tain temperature conditions so constant that the atmo- tials does cell extension occur with cell division to per-
sphere remaitis saturated yet no condensation of water tnit continuitig radicle growth. Even then, full itnbibition
on the seeds occurs. Machalica (1926) has reported that is not essential, there being a range of water potentials
germination is greater in a moist atmosphere with fluc- over which germination will occur.
tuating compared with constant tetnperature, and
fiuctuating temperature has also been shown to increase
seedling emergence from dry soil, probably through an (5) Differences between individual seeds
effect on the water vapour relatiotis of germitiation There now seems to be genetal agreement with the view
(Hegarty, 1975). Second, Christensen (1973) has pointed expressed by Hunter & Erickson (1952) that seeds must

reach a critical level of hydration to permit germination. hydration, althougli the beneficial effects tnay be
It does not, however, seetn to be equally generally ap- counteracted by damage, either gross or more subtle,
preciated that individual seeds eveti within a seed lot or by other responses on dehydration.
can hydrate to quite different levels before gennination In peas, during the initial stage of rapid hydration,
commences. the resistance of the seed to dehydration remains lugh,
In a detailed study of water uptake in calabrese seeds but subsequent uptake of water to the hydration level at
placed in contact with polyethylene glycol solution of germination causes a rapid decrease in dehydration
-0.75 MPa at 10C (Hegarty, utipublished) the indi- resistance, apparently associated with the accumulation
vidual seeds were weighed at frequent intervals for 35 of water in cytoplastnic vacuoles (McNair, 1966, quoted
days or utitil germitiation occurred. All seeds showed a by Sutchffe & Bryant, 1977). Dehydration of oat seeds
lag phase in hydration level after initial imbibition and following imbibition and prior to the first signs of visible
before the start of germination, and of 50 seeds used 32 germination has no effect on level or rate of gertnination
germinated between 11 and 35 days after the start of on subsequent rehydration, suggesting that the titne
imbibition and a further 15 seeds gertninated normally required for cotnpletion of germination by a population
on transfer to water after 35 days. Of the 32 seeds that of seeds depends upon the total time they are hydrated,
germinated at -0.75 MPa the mean water uptake just irrespective of whether or not this is interrupted by a
before germination relative to the original seed weight dry period (Akalehiywot & Bewley, 1977). However,
was 54%, with a range of from 44 to 66%, whereas the Berrie & Dretman (1971) considered that the effects of
tnean water uptake of the other 15 seeds after 35 days hydration before dehydration were only cumulative in
was 62% (significantly higher than 54%) with a range oat seeds if the hydration period was longer than a
of from 50 to 83%. Thus although the ranges of seed certain minitnum duration.
water uptake overlapped, the seeds that did not germi- Sen & Osborne (1974) found that getmination in
nate at 0.75 MPa bars were in general more hydrated, low-viability rye {Secale cereale L.) etnbryos was en-
not less, than those that did, and would only germinate hanced by a suitable hydratioti-dehydration treatment of
in an environment of higher water potential. Owen the etnbryo, and that there was a balance between the
(1952) also found that non-germinated seeds of wheat advancement of synthetic processes due to hydration
had higher moisture contetits than those that had and datnage to the earUest activated cells incurred
gertninated. during subsequent dehydration, where datnage was indi-
A different facet of this situation is expressed in the cated by the loss of ability to incorporate leucine
germination responses of seeds to moisture stress in (protein synthesis) or thymidine (DNA replication).
which, for a particular lot of seeds, germination is de- The hydration pretreattnent initially enhanced the
creased progressively over a range of decreasing water ability of the embryo to synthesise protein and RNA
potentials until no germination at all can occur. Relevant compared to the untreated controls, and the nortnal 9 h
to this is the statement of Crocker & Barton (1957) lag before the onset of DNA rephcation could be abol-
that low vigour seeds may need tnore water to germinate ished. Thus at least some of the changes occurring in the
compared to high vigour seeds, and it is also relevant initial stages of germination were stable to dehydration
that seed deterioration or loss of vigour is claimed to and subsequent storage. Support for this suggestion
lead to a diminution in the range of conditions in which comes frotn the earlier work of Marcus ct al. (1966) who
seeds can germinate (Abdul-Baki & Anderson, 1972). showed that desiccation did not destroy ribosomal ac-
Thus it is not the level of hydration per se that governs tivity in wheat embryos, and of Chen, Sarid & Katchal-
whether or not a seed will germinate. ski (1968) who also investigated hydration-dehydration
The work of Stiles (1948) on the distribution of treatments in wheat embryos. They found that rehydra-
water within seeds has already been mentioned and it tion for 24 h led to the transcription of new and active
would clearly be of considerable interest to know how mRNA in embryos that had been previously hydrated
reduced water potential affects the hydration levels of for 24 h but in those previously hydrated for 72 h only
the various seed structures, especially in individual trace atnounts of complementary RNA were transcribed,
seeds, and how it might also affect the inter-relation- and the transcribed RNA differed from the mRNA
ships between the embryo and its surrounding or en- fonned in normally germinating embryos. It was sug-
closed reserve materials. gested that on dehydration there was a breakdown of
DNA in 72 h etnbryos. DNA is actively duplicated and
transcribed in 72 h embryos and is relatively inactive
(6) The effects and timing of dehydration during the early stages of gennination. Thus it is pos-
Pre-germination hydration treatments can result in tnore sible that datnage on dehydration is associated with
rapid subsequent germination than in untreated seeds, DNA rephcation and RNA transcription.
although this may be partly due to more rapid imbibi- This more subtle datnage to the etnbryo should per-
tion resulting from a change in the physical structure of haps be distinguished from gross damage incurred when
the seed or seed coat (Kidd & West, 1919; Heydecker the growing embryo is dehydrated, although the two
et al., 1975; Heydecker & Coolbear, 1977). There is, may be related. The Russian work on seed 'hardening'
however, increasing evidence that the effect of physio- (May et ah, 1962) suggested that the latest stage at
logical development in the seed need not be lost on de- which seeds should be dried back was at the time of

radicle emergence frotn the seed (i.e. as cell extension The effects of dehydration on germination response
growth comtTiences, and possibly also coinciding 'with described above seem to fit very well the responses of
the start of cell division, c.f. Bertie & Drennan, 1971) over 40 grass, weed or woody species of Utah range
and Hubac (1962) has said that the ability of seeds to plants subjected to alternate wetting and drying cycles
survive dehydration is lost as hydration of the vacuoles (Griswold, 1936). Griswold concluded that prior to ger-
of the embryo eommenees. Ho'wever, Carcellar & Sori- mination, interruption of the moisture supply may
ano (1972) dried back wheat seeds just after germination arrest the development or growth of the embryo which
without deleterious effeets on subsequent germination, precedes germination until sufficient moisture is sup-
and even if this is done repeatedly, viability may be re- plied again, whereas dehydration may alter the proper-
tained in some seeds over several cycles of hydration ties of the seed coat, may produce secondary dormancy
and dehydration, with cereals being more resistant than or, if at a critical stage, may injure the embryo beyond
peas (Widstoe, 1916). Seedlings at an even more ad- the point of recovery.
vanced stage of germination can be dehydrated without One final effect that may be of relevance is the
causing death, although growth may be resumed from phenomenon of vernalization which is induced in seeds
secondary initials (Milthorpe, 1950; Hegarty, 1977a). held fully or partly imbibed in the cold (Napp-Zinn,
It is interesting that isolated embryos of low-viability 1961). The phenomenon is apparently reversible,
rye when at too advanced a stage before dehydration althougli the loss of the effect when it does occur may
also initiated growth from secondary initials (Sen & be due to too high a temperature being maintained
Osborne, 1974). during dehydration rather than to the process of de-
In oats, Berrie & Drennan (1971) found that protease hydration itself (Schwabe, 1971). There seems to be
activity was higher in treated seeds on subsequent im- no physiological advantage in using the Russian method
bibition than untreated seeds, that this response was vvhicli prevents germination during the treatment, tliis
stable in stored seeds and that the effect of successive being considered to be simply more convenient for
hydration-dehydration cycles was cumulative only if the subsequent handling of the seed (Purvis, 1961).
prior imbibitions were of substantial duration. In crested Thus some at least and possibly many of the physio-
wheat grass (Agropyron desertotttm (Fisch. ex Link) logical developments associated with the preparation of
Schult), Wilson (1971, 1973) found that incubation of the seed for gerrnination appear to be stable to de-
seeds in vapour over solutions with water potentials hydration if they are not counteracted either by cell or
down to -4.0 MPa, or seeds placed in soil and recovered organ damage in the embryo (which may coincide with
at various times during the gertnination process, showed the initiation of cell elongation or cell division) or by
an increased subsequent rate of germination at low tetn- some form of secondary dormancy. It seems possible
perature, with the increase in a-amylase activity being a that an important difference between pretreatment in a
good indicator of the hastening effect of the treatment. restricted quantity and in a free supply of water eould
In a series of eight cycles of hydration-dehydration, a- be the extent to which substrates can accumulate or
amylase activity was successively accutnulated without system activity can be enhanced in the seed for use
loss on dehydration. In contrast, seeds sown on the soil during subsequent germination.
surface showed increase ATP levels after rainfall but Also relevant in this Section is the contrast made by
these levels fell again under severe tnoisture stress Thomas (1972) between the process of dehydration in
(Nelson, Wilson & Goebel, 1970). maturing seeds and rehydration after maturation. Some
Futther evidence for either accumulation of sub- of the marked changes in cell organisation wliich occur
strates or enhancement of activity of enzymes or systems as the seed matures are direct expressions of changes in
in seeds prevented from germination by osmotic stress, the state of cell membranes atid may be related to bio-
comes from the subsequent inereases in respiration rate chemical changes in dehydrating seeds. The maturing
after dehydration and imbibition compared to non- seed is subjected to water stress probably severe enough
treated seeds. This has been reported for pepper (Wood- to contribute to marked changes in respiratory activity
stock, 1969; Malnassy, 1971), tomato (Woodstock, and enzyme synthesis, and Thomas accepts the sug-
1969; Koehler, 1967), and Dactylis glomerata (Haight & gestion of Klein & PoUock (1968) that ehanges in the
Grabe, 1972). This effect might resemble the 'stored physiological activity of the tnaturing seed are adapta-
growth effect' described by Ray (1961) in which oat tions which render the cells resistant to desiccation. In
coleoptile growth is reduced by transfer to osmotica but addition, he poses the question whether water stress
then surges forward itiitially at a much greater rate on might alter growth regulator levels in maturing seeds (as
transfer to water. This is apparently not simply a physi- it does in leaves), with an implied control over metabol-
cal effect associated with osmotic properties because the ism. He also points out that the proteins synthesized by
stored growth effect can be inhibited by KCN. maturing seeds are very different to those made by
One further response resulting in the negating of germirtating seeds, and presumably the mRNA's must be
beneficial effects of seed 'priming' of celery has been different in tlie two cases. It follows that at some stage
described by Gibbins & Heydecker (1977) in which a during seed development there must be destruction of
germination inhibitor apparently accumulates in the the old messetiger and a synthesis (and possibly storage)
seed on dehydration, causing a form of secondary of the new, and it is possible that desiccation, by
dormancy. directly affecting cell nuclear volume and the osmotic

and ionic environment of the chromatin, causes the either set of circumstances may give a survival advantage
ordered openitig up or closing down of genetic areas. to seedlings (Wilson, 1971; Wilson, Wotidercheck &
If Thomas's hypothesis is correct, then, by analogy Goebel, 1974). In less extreme environtnents, tnoisture
with the maturing seed, it is possible first that at sotne stress can delay seed germination in the soil and can
stage during the germination process or during the seed result in a double fiush of seedlings after subsequent
priming treattnent the etnbryo may lose its adaptation rainfall. A laboratory experiment with calabrese and
to desiccation and hence suffer damage if subjected to carrot seeds suggested that pre-etnergence losses in the
dehydration, and second, that desiccation during the particular conditions used were due to seedling dehydra-
gertnination process may itself cause a change in genetic tion rather than to datnage of the seeds (Hegarty,
response to subsequent rehydration. 1977b), although the precise degree of seed hydration
One possible site of damage during desiccation is the may be very important, especially with respect to the
cell metnbrane and, in his review on freezitig injury in interaction of the seed with seed or soil pathogens
cells, Merytnan (1974) argues persuasively that solute (Wallace, 1960).
concentration damage may not be the result of de-
naturation by the concentration of any specific solute,
{1.3) Dormartcy
but appears to be related to the osmotic reduction of
cell volume, with the possibility that the cell, and par- Several types of dortnancy yield situations in which
ticularly the tnembrane, is damaged mechanically if a there is active seed metabolism yet no gertnination. In
cell decreases in size below a tninimum critical volume. some species, after-ripenitig must take place in itnbibed
He suggests that there is a mechanical resistance to seeds before specific dormancy-breaking treatments
volume change in cells associated with the metnbrane leading to germitiation are started. In such seeds, ana-
and, as a result, the internal hydrostatic pressure of the tomical and morphological changes tnay occur (Mayer &
cell matrix contained within the plasmaletnma can fall Poljakoff-Mayber, 1975). Detailed studies in ash {Fraxi-
below attnospheric pressure, thus creating a pressure tws excelsior L.) have revealed tliat althougli no cell
gradient across the membrane. This, in turn, can lead division occurs during the after-ripening phase, there are
to an abrupt change in permeability of the metnbrane extensive food cotiversions, wall development and the
(the site of pritnary damage), permitting an influx of appearance and proliferation of cellular organelles
extracellular solutes and loss of stability of the metn- duritig this phase of deep dormancy (Villiers, 1971).
brane macromolecules, thus pertnitting further In dormant seeds sliortly after imbibition, rates of
degradative events leading to irreversible change. respiration as high as or approaching those of non-
Although this hypothesis is proposed to account for dormant seeds have been found in wheat (Miyamoto,
the effects of freezing injury in cells, with the etnphasis Tolbert & Everson, 1961 ;Ching& Foote, 1961), in wild
being on animal cells, it clearly has itnpHcations for the oat {Avena fatua L.) (Chen & Varner, 1970) and in
process of dehydration in seeds after hydration, and, if lettuce (Foutitain & Bewley, 1976), suggesting that
cell shrinkage were differentially affected, could offer seeds are active at that time though they cannot gertni-
an explanation for differences in response due to dif- nate. In barley and rice {Oryza sativa L.), initial rates of
ferent rates of drying. respiration were even higher in dormant thati in non-
dortnatit seeds, but the pathways of respiration were
apparently different in the two states (Major & Roberts,
(7) Analogous situations 1968).
Seeds can survive, imbibed, in the soil for long
(7.1) Salt marsh species periods (Crocker, 1916) yet respiration rates of such
The closest analogy to the seed 'priming' treatment seeds can be appreciable (Barton, 1945), adding support
occurs with seeds released frotn salt tnarsh plants into to the theory that seeds in soil tnay eventually die due
sea water (Chapman, 1960; Boorman, 1968). Seeds of to exhaustion of food reserves (Crocker, 1916). The
the tnajority of halophytes, although apparently resis- rate of respiration found for non-dormant calabrese
tant to the damaging effects of NaCl, germinate best in seeds held under moisture stress in polyethylene glycol
fresh water conditions. If first held in sea water, they solution seemed to be cotnpatible with survival for a
germinate very rapidly on transfer to fresh water. Boor- long period before the seeds' reserves would have been
man considered that the early stages of the germination exhausted (Hegarty, 1977a).
process could take place in salt water and only the final Itnbibed seeds of lettuce held under induced thertno-
stages preceding germination were inhibited. dortnancy (30C) in Petri dishes maititained their
viability over 105 days, but viabiHty was even better
maintained in seeds stored at 30C in a RH of from 85
(7.2) Arid environments to 90% (Toole & Toole, 1953). In an extension ofthis
Successive cycles of wetting and drying can occur in work (Villiers, 1974) seed viability in lettuce and ash
arid environments and are likely to permit developtnent {Fraxinus americana L) was maintained over 12 months
of the embryo, resulting in more rapid germination in seeds kept fully imbibed, but was cotnpletely lost in
either when substantial rainfall occurs, or at low tem- 2 months (lettuce) and 3 months (ash) in seeds equili-
peratures in the spring. Rapidity of establishment in brated at 100% RM and then stored. Chromosotne

aberrations increased sharply in the vapour-equilibrated cell division and elongation on the one hand, and the
seeds at increasing moisture contents but were absent in initiation of cell elongation on the other, may, if appli-
the fully imbibed seeds, indicating an ability of tlie cable in seeds, be of fundamental importance in the
dormant seeds to prevent storage deterioration. The control of germination. If the water potential threshold
water content of the lettuce seeds equilibrated at 100% for the initiation of cell elongation is higher than that
RH was only 14% (dry weight or wet weight not indi- for other processes of cell development (i.e. if the initia-
cated) and Villiers speculated that a higher moisture con- tion of elongation is the process most sensitive to
tent would be needed before macromolecular repair moisture stress) then at the appropriate water potential
and membrane synthesis could occur at a normal rate. seed development miglit be permitted but not active cell
Dormancy can also be due to physical constraint of elongation, and the initiation of cell elongation would
the embryo by the seed coverings (Crocker, 1916; be the block in the germination process that allows seeds
Mayer & Poljakoff-Mayber, 1975). In this situation to be brouglit to the 'brink' of germination but no
embryo activation clearly occurs and the constraint further.
may be simply due to the strength of the seed coat.
Indeed, pressure applied to the soil surface can, if suffi-
cient, prevent germination in non-dormant seeds (Collis- (8) 'False' germination
George & Williams, 1968). Up to this point it has been loosely assumed that germi-
It has been implied that dormancy is an inactive nation, identified as radicle growth from the seed,
resting phase (Toole & Toole, 1953; Yemm, 1965). represents a continuing phase associated with concurring
Haber & Luippold (1960b) suggest that such an implica- cell division and cell elongation. Rogan & Simon (1975)
tion is quite inappropriate to describe the physiological have described an initial phase of slow elongation of
state of seeds prevented from germinating by with- the radicle immediately after imbibition before a more
holding morphogenic stimuli. It is not generally related, rapid phase commences and before mitosis is detected.
either, to a significant depression of overall metabolism This phase of growth is therefore presumably associated
or to a block preventing cell division. Instead, they only with the elongation of existing cells in the embryo,
suggest tliat it is a subtle block that specifically prevents and Haber & Luippold (1960a) concluded that rootlet
the initiation of cellular expansion. In many ways this protrusion in lettuce is a result of cell elongation, with
situation seems to be remarkably similar to seeds held at cell division playing no part. If conditions are such that
the brink of germination by moisture stress, as in the some cell elongation is permitted but no cell division,
seed 'priming' treatment. than a 'false' germination may result. This has indeed
been described in, for instance, sorghum (Sorghutti
vulgare Pers.) (Nutile & Woodstock, 1967) and in some
(7.4) The physiology of growing root eells grasses (Watt, 1974). In the latter case, radicle protru-
The subject of the transformation of meristematie into sion witliout further growth was a response to low water
elongating cells in roots has been reviewed by Obrou- potential, occurring between 0.5 and I.O MPa, and
cheva (1975). Both transformation and elongation are appeared to be a resting phase before growth continued
accompanied by acceleration of all metabolic processes. in some seeds at the same water potential, or in all
The content per cell of numerous compounds is much seeds on transfer to free water. In one species in
higher in elongating than in meristematie cells, both in particular, these 'hydropedetic' seeds were resistant to
the case of compounds of primary metabolism as well as dehydration injury and germinated readily on
the end products. Most compounds of primary metabol- rehydration in free water.
ism are osmotically active and tlie increases in their levels Hadas & Stibbe (1973), considering tliis phenomenon
in elongating cells favour decreased solute potential and in chickpea (Cicer arietitmtn L.) and maize seeds, suggest
increased water uptake. These changes are accompanied that there exist critical hydration levels within the seed
by parallel increases in enzyme activity and in respira- which will permit various growth processes. The first
tion, althougli the balance between the predominant critical level permits cell elongation whereas the second
respiratory pathway through glycolysis and the Krebs permits cell division in the radicle.
cycle on the one hand, and through the hexose mono- Whilst it is probably the initiation of cell elongation
phosphate pathway on t)ie other, remain constant. Thus that is sensitive to water stress in this instance, the idea
elongating cells represent sites of active metabolism of of differential sensitivity of different growth processes
a type that, as in meristematie cells, is dependent upon to water stress is supported by the evidence that the
the normal course of protein and nucleic acid syntheses threshold water potential for shoot growth is higlier
and is sensitive to inhibitors of those processes. How- than that for root growth (Dasberg, 1971; Gray &
ever, cell elongation does not necessarily follow the Steckel, 1976). 'Sprouted' but non-emerged seeds were
completion of cell divisions in the meristem, and found by Doneen &, MacGillivray (1943) in dry soils,
although the mechanism governing the process of trans- and these may represent 'false' germination, or radicle
formation itself and the initiation of elongation is not growth without shoot growth as described by Dasberg
known, it has been found to be insensitive to inhibitors (1971). Restricted radicle growth without shoot growth
of protein and nucleic acid syntheses. is also a phenomenon of low temperature (Kotowski,
The dependence on different metabolic processes of 1926) and this would coincide witli the finding of Haber

& Luippold (1960a) for lettuce that at low temperature reduction of seedling mortality. He considered that
mitosis is restricted more than cell elongation. recent work on the initiation of germination has concen-
trated on dormancy mechanisms, both innate (primary)
and imposed (secondary). In general, dormancy is a
(9) Does the osmotic inhibition of germination resemble means by wliich seeds avoid germinating at a time when
a dormancy mechanism? seedlings would be placed in high-stress conditions.
Evidence is accumulating that germination that is just The failure to germinate under moisture stress as des-
inhibited by osmotica such as polyetliylene glycol or by cribed in the previous section would constitute a simple
NaCl can be restored to a greater or lesser extent by a example of an imposed dormancy mechanism which in
stimulus given to the seed directly or through the germina- many species (lettuce may be an exceptionKalm,
tion medium. The evidence is particularly clear in the case 1960) is apparently readily reversible in an improved
of lettuce, in which osmotic inhibitionhas been overcome water supply. This, together with the differential sen-
by gibberellic acid added to the germination medium sitivities of radicle and shoot growth discussed in Section
(Kahn, 1960), by red light (Kahn, Goss & Smith, 1957; 8, would have obvious survival advantages in dry condi-
Schiebe & Lang, 1965; Nabors & Lang, 1971) and by tions. Certainly, the ability of seeds to germinate at low
growth regulators applied to the seed (Odegbaro & water potential can lead to establishment failures if dry
Smith, 1969; Kaufmann & Ross, 1970; Braun & Khan, weather ensues, whereas seeds with more exacting re-
1976; Khan & Knypl, 1977). quirements for germination (often species native to semi-
Whilst lettuce may not be typical in its response arid areas) can establish more successfully (McGinnies,
there are, in addition, other reports of a similar nature in I960; McWilliam & Phillips, 1971; Mott, 1974; Watt,
other species. Osmotic inhibition in celery was overcome 1974).
by growth regulators applied to the seed (Klian & The first stage of imbibition is the result of the
Knypl, 1977) whilst partial release of salt effects using physical properties of the seed (Mayer & Poljakoff-
growth regulators has been described for wheat (Chaud- Mayber, 1975), and Yamamoto (1964) has proposed
huri & Wiebe, 1968; Idris & Aslam, 1975), pea (;Uprety that the retention of absorbed water may be an adapta-
& Sarin, 1973) and Spergularia media (L.) C. Presl. tion to xeric conditions. It might equally well be argued,
(Ungar & Binet, 1975). In wheat, presoaks in some however, that rapid dehydration could prevent germina-
growth regulators were more effective than water alone tion of seeds which, had they germinated, would have
in overcoming osmotic inhibition during subsequent exposed the seedlings to a higli moisture stress environ-
gennination (Darra et al., 1973). Finally, gennination in ment (Mott, 1974).
the dark was higher than in the light in osmotically An alternative strategy promoting seedling survival in
stressed seeds of cucumber (Cucumis sativus L.), radish drying conditions would be the ability for seedlings
and sunflower (Helianthus annuus L.) (MeDonough, after germination to grow at water potentials lower than
1967). those permitting germination. Some evidence for this is
Although in many cases only a portion of the seed provided by the results of MeDonough (1975) who
population was released from the osmotic inhibition, showed that seeds removed from a water supply after
nevertheless there is a phenomenon common to all initial imbibition could continue to germinate, and that
these reports. Seeds were stimulated to germinate in equilibrium water potentials were eventually attained
conditions in which, without the stimulus, they would which were lower than those that could have permitted
not have germinated even though the conditions of germination.
water supply were the same. Thus there is a strong sug-
gestion that the failure to germinate under moisture
(11) Membrane effects
stress, in some species at least, represents a positive
response to the environmental conditions and is, in fact,
(11.1) Membrane integrity
a form of secondary or induced dormancy. In many
respects this phenomenon resembles the responses of As mentioned in Section 4.2, seeds soaked in water
some seeds to high temperature in which germination imbibe, and can lose a wide range of solutes. The leach-
inhibition can be overcome by treating the seeds directly ing of solutes into the soak water has been used as a
with growth regulators (Biddington & Thomas, 1976; seed vigour test in large-seeded legumes such as peas and
Heydecker & Joshua, 1976; Dunlap & Morgan, 1977). French beans (Matthews & Bradnock. 1968) and in
field beans (Vicia faba L.) (Hegarty, 1977c). In peas,
seeds with dead tissue on the abaxial surfaces of the
(10) Ecological relevance cotyledons showed the highest level of leaching but the
In a recent review Matthews (1976) proposed that seed large range of levels found among seed lots resulted from
and seedling adaptations were ecologically significant differences in leaching from live tissue (Matthews &
because they tended to minimize post-germination Rogerson, 1976). Whilst increased solute leakage may be
seedling losses. He differentiated between adaptations partly due to induced cotyledon damage in these large
which influence the initiation of germination and those seeds during imbibition (Pollock, Roos &Manalo, 1969;
which influence the chances of seedling survival, but Rowland & Gusta, 1977), membrane permeability ap-
recognized both as being means to the same end, the pears to be the main property governing the extent to

which leakage will occur. This subject has been exten- though not related to germination in particular, and the
sively reviewed by Simon (1974) who considers that effect on this of growtli regulators acting via the mem-
leakage occurs until the phospholipid membranes of the brane, have been reviewed by several contributors in the
cells become intact, with this occurring most rapidly in volume by Marre & Ciferri (1977). In addition, Cocucci
the outer layer of cells. Loss of seed vigour associated & Cocucci (1977) point out that fusicoccin appears to
with an inciease in seed leachate is put down to a de- be especially effective in overcoming germination
terioration of the lipid membrane. In addition, the inliibition in lettuce resulting from moisture stress or
imbibitional cliilling injury found in seeds such as maize, liigh temperature (Braun & Khan, 1976), thus again
cotton, lima bean (Phaseolus Iwtatus L), French bean, suggesting control over transport relationships.
soybean and sorglium, which can be prevented if the The potential importance and involvement of the
seed moisture content is raised before chilling (Cohn & cell membrane has been further indicated by the possi-
Obendorf, 1976), if the imbibition takes place for a bility that pressure itself may control transport processes
short period at a higiier temperatuie before chilling in the plasmalemma and tonoplast. Coster, Steudle &
(Pollock & Toole, 1966), or, in cotton, if the seeds are Zimmermann (1976) suggest that the cell membrane is
given a hydration-dehydration treatment at higli tem- compressible by both mechanical and electrical forces,
perature before chilling (Thomas & Christiansen, 1971), and that significant changes in the tliickness of the
is also thought to be associated with membrane proper- membrane can occur as a result of direct compression
ties (Simon, 1974). due to liigli turgor pressure, indirect effects due to
Bramlage, Leopold & Parrish (1977) have reported stretching of the cell wall, and the stresses induced by
that in soybean, humidifying the embryos to 25-30% the electric field in the membrane. They suggest that
moisture (dry or wet weight basis not indicated) reduced changes in membrane thickness could provide the
leakage and overcame cotyledon cracking during imbi- pressure-transducing mechanism required for osmo-
bition, but did not eliminate continued leakage at low regulation and growth. Hence the rate of pumping of
temperature or imbibitional chilling injury, whereas in ions (and thus the solute potential of the cell sap) could
embryos humidified to 35-50% moisture before imbibi- be directly controlled by the turgor pressure.
tion, leakage at low temperature and imbibitional
chilling injury were eliminated.
(12) Germination control
From work on peas, Simon & Wiebe (1975) propose
that embryos with a moisture content of 43-54% (water There is a major problem in bringing together informa-
potentials of 8 to 5 MPa) will show httle or no tion from widely different species involving different
leakage during imbibition. The degree of hydration of seed structures, with the accompanying different re-
the membrane at these water potentials is not known, lationships between the embryo and its enclosed or
but it is suggested that matric forces in the cells are surrounding reserve material, different seed constituents
reduced to such an extent that water is available to stabi- and different dormancy types and levels, in which ger-
lise the membrane bilayer. This level of water availability mination control mechanisms may be entirely different.
is sufficient to permit the formation of ice crystals on A large amount of work has been carried out on the
freezing. physiology of lettuce seeds, yet the relevance of this
work to seeds in general is not at all clear. From tWs
sort of work, however, has developed the concept of
(11.2) Other membrane properties germination control resulting from a balance of growth
Leakage of salts from seeds has also been reported for regulators within the seed (Khan, 1975), with germina-
radish (Cocucci & Cocucci, 1977) but in this case the tion probably operating through a matrix of interacting
leakage was followed by active salt uptake, at least in metabolic pathways and with growth regulators influ-
the case of K'^. Addition of abscisic acid (which inliibits encing the enzyme activities within this matrix (Hey-
radish germination) to the medium enhanced the salt decker & Coolbear, 1977). Added to this is the problem
loss whereas fusicoccin (which stimulates the rate of that release of germination inhibition by stimuli (as
germination) promoted rapid, active K'^ uptake. The opposed to the leaching of an inhibitor) may act by
authors suggest that activation at the cell membrane different means. In the case of lettuce, for instance, it
level of some metabolically-dependent mechanism of has been suggested (Schiebe & Lang, 1965) that the
K* uptake probably linked to H"^ release is associated dormancy caused by embryo constraint within the endo-
with germination in radish seeds. Fusicoccin probably sperm can be broken by irradiation in red light, inducing
enhances K* uptake capacity whereas abscisic acid in- greater embryo thrust, or by gibberellin which possibly
liibits it, and germinatioii could be linked with H* acts by weakening the endosperm.
weakening of the cell wall and/or K"^ uptake decreasing There are, however, some significant features that
the solute potential of the cell, both mechanisms en- can be extracted from the data wliich may point to an
couraging water uptake. understanding of germination control and the effects of
A general role of membranes in the control of germi- seed hydration treatments.
nation was suggested by Mayer & Sliain (1974) on evi- (1) In the work just mentioned (Schiebe & Lang,
dence from several sources. The involvement of the 1965) it was found that red hght caused increased em-
membrane in the specific control of solute transport. bryo growth of lettuce seeds in mannitol solution. This

work has now been taken further to show that phyto- are at least partially responsible for determining the
chrome stimulation of lettuce axes held in polyethylene success or failure of germination.
glycol solution can result in a decrease of solute poten- (7) The fact that in a population of seeds different
tial within the embryo of 0.1-0.2 MPa, and a change in numbers germinate over a range of water potentials,
pressure potential of a similar order such that in total and that pre-soaking treatments can alter these responses,
the embryo overcomes the constraint of the endosperm imphes that seeds in a population have a range of
(Carpita & Nabors, 1977). According to Black (1969) threshold levels for initiation of germination and that
seeds must be undergoing active respiration in order for the threshold levels can be altered. This could be inter-
phytochrome to act. The point is that given certain preted as a membrane effect (e.g. allowing hydration or
external conditions, osmoregulation within the seed was even repair under favourable conditions) but in any case
clearly affected by the external stimulus. it is evidence that events occurring later in germination
(2) Pressure can affect cell membrane properties and, are dependent upon events occurring earlier in the germi-
through this, osmoregulation (Coster et al., 1976). nation process.
Changes in water potential could well affect cell mem- (8) Some seeds can imbibe and be held at low tem-
brane properties either by acting directly on the mem- peratures for very long periods yet still germinate nor-
brane, or indirectly by affecting the pressure relations of mally on transfer to favourable conditions (Harrington,
the cell. 1962). In a species such as carrot, hydration treatments
(3) Growth regulators could simOarly act directly on at temperatures favourable for germination can help to
ion transport processes in the cell membrane (as sug- overcome low-temperature sensitivity (Hegarty, 1970)
gested by Marre (1977) for fusicoccin), or indirectly and presumably in these cases membrane effects as
through cell metabolism but ultimately affecting trans- described in paragraph 6 above are not involved. Austin
port processes in the membrane (e.g. weakening of the et al. (1969) have suggested that in such cases hydration
cell wall). Braun & Khan (1976) point out that effects treatment may allow the embryos of some seeds to
of water stress have been associated with alterations in 'after-ripen' and in this case it may be chemical or mor-
the levels of endogenous growth regulators and, in seeds, phological, or possibly membrane, modifications that
exogenously applied growth regulators can sometimes permit some seeds to germinate at low temperature that
break dormancy, including that induced by water stress. would not have germinated without the hydration
Clearly, this action could be through an effect on the treatment.
cell membrane and solute transport. (9) Germination itself must be associated with the
(4) Some cell division may apparently occur in the osmotic regulation of cells in the radicle. Such osmotic
embryo at water potentials lower than those that permit regulation can apparently be influenced in a number of
the co-ordinated division and extension associated with ways in seeds. It seems likely that many, if not most of
continuing radicle growth. If the situation described in the processes that occur in seeds supplied with water
roots by Obroucheva (1975) can be applied to the ger- sufficient to germinate also occur in seeds held under
minating seed, then it may well be the process of the moisture stress such as that of the seed 'priming' treat-
initiation of elongation, which apparently has quite ment (though perhaps more slowly), the main differ-
distinct metabolic properties, which is involved in the ences being associated with the process that induces cell
control of radicle emergence and its continuing growth. elongation. It may well be significant that this process
This process is presumably concemed with osmoregula- could be under a metabolic control that apparently does
tion within the elongating cell. The work of Fujisawa not infiuence other processes within the seed associated
(1966) who showed that phase C growth of radish axes with preparation for germination (i.e. phases A and B of
could be inhibited without interfering with phase B germination).
respiration, miglit also suggest that it was the initiation (10) There is a strong resemblance between the
of cell elongation that was being inhibited. 'subtle' block to germination described by Haber &
(5) The proposal that the transformation of cells to Luippold (1960b) for some types of dormancy and the
the elongating stage has a higher water potential threshold block to germination of seeds held under moisture stress.
than embryo development or growth, implies as a conse-
quence that germination once initiated could proceed at
(13) Conclusions
lower water potentials. This is supported by the findings
of MeDonougli (1975, 1976) that germinated seedlings Initial seed imbibition is apparently a physical process.
do develop low water potentials. Seeds in contact with a free water supply take up more
(6) In some cases, imbibition under unfavourable con- water than they need to germinate, though the absolute
ditions affects subsequent germination, presumably quantity taken up will depend on the seed structure and
through an effect on the hydration of membranes constituents, on the temperature and possibly on other
(Simon, 1974) and initial, even partial, hydration under properties of the external environment. At lower levels
more favourable conditions can overcome this problem. of water potential seeds imbibe less and germinate more
Seed ageing results in membrane deterioration and a slowly until eventually germination is prevented. This
reduction in the range of conditions in which seeds will varies between individual seeds within a seed lot, be-
germinate, and it is possible that in low-vigour seeds of tween seed lots of the same species, and between species.
this category, membrane hydration or integrity effects Even when seeds are prevented from germination by

the water potential of the external etivironment, it is activation or repair processes counteract or replace the
abundantly clear that there is a range of water potential mechanisms of seed deterioration. A limited atnount of
within which many, if not all, of the processes leading to work suggests that membrane integrity is not achieved
normal germinatioji up to a particular point can occur, until water potentials of 8 to S MPa are attained, not
and that substrates for future growth may accumulate, very much less than the levels (down to - 2 to - 4 MPa)
or systetn activity may be enhanced. Several aspects of usitig osmotica or vapour treattnents at which known
the environment appear to affect the precise extent to benefits to seeds have been reported. It is possible that
which all the seeds within a seed lot may be brought to seed activation or repair mechanisms cannot operate
the brink of germination. Seeds unable to germinate for either effectively or at all until tnembrane integrity has
other reasons (various types of dormancy) have also been attained, and that deterioration mechanisms domi-
been shown to be metabohcally active, and sotne forms nate or operate alone at hydration levels below these
of dortTiancy show similarities to the inhibition of germi- water potentials (8 to 5 MPa). Tliese levels are in
nation by moisture stress. equilibrium with very higli RHs (95% +) and the prob-
Dehydration before germination tnay suspend activity letns associated with maintaining accurately such RHs
and development at the level just prior to dehydration, could well explain at least some of the conflicting
but may also be associated with gross or more stibtle results obtained with seeds held in so-called saturated
forms of embryo damage (includitig damage to tlie cell atmospheres.
membrane), or with the induction of fotms of secondary The likelihood of finding common trtechanisms of
dormancy. gertnination cotitrol and seed activation or deterioration
One can speculate that the process in the embryo under moisture stress seems in many ways highly un-
most sensitive to moisture stress is the initiation of cell likely because it is diversity that encourages survival.
elongation. This is likely to be associated with osmo- Nevertheless, whilst speculative, some of the suggestions
regulation within the cells. It appears that osmoregula- above do offer links between the findings of many ex-
tion is sensitive to various factors within the embryo perimenters working with quite different objectives in
which tnay act independently or through a common mind, and several lines of work suggest that the cell
pathway, and mucli evidence suggests that the cell tnetnbratie tnay be futidatnentally involved. Tliis hy-
metnbrane is actively involved. Osmoregulation is sensi- bridisation of ideas may stimulate a number of critical
tive to external stimuli and this suggests that the failure experiments to be performed which, at very least, should
to germinate under moisture stress can, in some cases at yield information on the physiology of incompletely
least, be a positive response to the environment, and can hydrated seeds, a field in which very little information is
be classed as a form of secondary dormancy. In roots, currently available.
the mechanism governing the initiation of cell elotiga-
tion has metabolic properties which are distinct from
the processes of cell division and cell elongation, and, if Acknowledgments
applicable in seeds, this could give the advantage of I am indebted to Dr John Raven of the Department of
allowing normal embryo development to occur in Biological Science, University of Dundee, for helpful
drought-stressed seeds whilst preventing germination. discussion during the preparation of tliis paper, including
The phenomenon of physiological development of the indication of a nutnber of relevant references, and to
the embryo without germination occurs in natural habi- Dr Peter Waister of the Scottish Horticultural Research
tats and, under some circumstances at least, may lead to Institute for useful comments on the content and
enhanced seedling sumval. Conversely, sowing pre- construction of the tnanuscript.
germinated seeds may place the seeds seriously at risk in
the environment by removing gertnination control from References
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