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
for the improvement of level, rate and uniformity of
gennination or field emergence has revealed how little
is Imown of the physiology of getmination control
under water stress. This review surveys briefly the re-
sponses of seeds held at different hydration levels, from
normal germination in a free water supply to seed
activation without germination under slight moisture
stress, seed deterioration at greater moisture stress and
the damage that can be caused to seeds in very dry con-
ditions, as well as the responses to subsequent dehydra-
tion. Inhibition of germination, though not of seed
activation, at certain levels of water stress is likened to
various forms of dormancy, and the mechanism govern-
ing the initiation of cell elongation is suggested as the
possible key to control over germination. Several lines
of evidence on cell membrane integrity and action, and
their responses to external factors, point to tlie role that
the membrane may play in cell elongation (and hence
germination) eontrol, and membrane integrity may
also be associated with the transition between seed
deterioration at one hydration level and seed activation
and repair at slightly higher hydration levels. Seed activa-
tion witliout germination is also considered in an eco-
logical context.
(1) Introduction
There has recently been a considerable upsurge of in-
terest in the use of pre-sowing seed treatments involving
the full or partial hydration of seeds, sometimes fol-
lowed by subsequent dehydration, which result in an
increase in the rate, uniformity or level of seed germina-
tion.* These seed treatments for improved performance
have been described, amongst many other kinds of seed
treatment, by Heydecker & Coolbear (1977). That the
treatments have substantial effects on the seed germina-
* In this paper 'seed' is used to describe all types of propagule,
'germination' is used to describe the emergence of the radiele
from the seed, and the 'germination process' is that series of
events culminating in germination itself.
Correspondence: Or T. W. Hegariy, Scottish Horticultural
Researeh Institute, Invergowrie, Dundee.
0140-7791/78/0600-0101$02.00
1978 Blaekwell Seientific Publications
tion process tliere is no doubt; the mechanisms of the
effect are, on the other hand, far from clear.
In addition, seed responses at different hydration
levels can vary widely, from germination on the one
hand to deterioration and death on the other, whilst
some seeds can remain dormant and viable in the soil
for many years while fully hydrated. It is the intention
in this review to consider many of these responses to-
gether to see whether it is possible to build a more
comprehensive picture of the relation between seed
physiology and the degree of seed hydration. Of particu-
lar interest are the precise responses of seeds to differing
levels of hydration; the effects of subsequent dehydra-
tion ; analogous or related situations that may yield rele-
vant infortnation on control mechanisms of seed
germination under moisture stress; and whether non-
germination under moisture stress is an active or passive
response to the external environment.
This is not intended to be a comprehensive literature
review. Rather, I have used, where possible, references to
general treatises or reviews which themselves include
many of the references to original work; references to
specific pieces of work are used where no review exists
or where specific poitits are being tnade. Consideration
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.
(2) Historical perspective of hydration treatments
Much of the work of the last century was reviewed by
Kidd & West (1919). They concluded from the results
published that seeds swollen in water and sown still in
the moist condition germinated more rapidly than un-
treated seeds; tliat seeds soaked in a 'minimum' amount
of water and afterwards dried slowly at ordinaty tem-
peratures imbibed water and developed more rapidly
when again allowed to take up water and germinate,
than untreated seeds; but that seeds dried rapidly after
initial soaking germinated tnore slowly than untreated
seeds. Particular reference was made to the conditions
of seed soaking such as the quantity of water, duration
and temperature of the treatment, and to the apparent
lack of effect of salts added to the soaking solutions.
Their own experiments (Kidd & West, 1918) indicated a
beneficial response to soaking in some cases but deleteri-
101
102 T. W. HEGARTY
ous effects in others. Chippindale (1934) investigated
the effects of seed soaking on grasses, and mentioned
that the practice was formerly carried out frequently by
farmers, but not in 'modern' agriculture. He concluded
that the effects found could not be generalized over
species, even when closely related.
A move towards making a more defined use of seed
hydration treatments was made by Le\att & Hamm
(1943) who were concerned to ftnd a way to 'mature'
seeds of Taraxacum kok-saghyz Rodin, before sowing.
They speculated that it should be possible to increase
the moisture content of the seeds sufficiently to pennit
the necessary physiological changes, but insufftciently to
permit gertnitiation, and then dry back the seeds so that
they could be sown in the usual way. Accordingly, they
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
solution (e. -2.7 MPa) or 0.5 M KNO3 solution (c. -2.0
All the methods hastened subsequent germitiation, the
optimum treatment incorporating molar (M) dextrose
solution (c. -2.7 MPa) or M/2 KNOa solution (c. 2.0
MPa), with optimum duration dependent upon the tetn-
perature. There was no loss of benefit in dehydrated
seeds stored for 4 weeks.
To a considerable extent this work seems to have re-
mained utirecognized, and only 20 or so years later,
possibly after the publication of a review of Russian
work on seed 'hardening' (May, Milthorpe & Milthorpe,
1962) was there an increase in interest in the use of
pre-sowing treattnents using a restricted quantity of
water. The Russian work was concerned with inducing
drouglit tolerance during subsequent plant growth, and
consisted essentially of variations on the use of one or
more cycles of seed hydration in a restricted quantity of
water followed by dehydration after a specified time. It
was considered that as germination proceeded the resis-
tance of the embryo to dehydration decreased and the
likelihood of permanent injury from drying increased
whereas the degree of 'hardening' induced was likely to
be greater the more advanced the embryo at the time of
dehydration.
Meydecker (1974), in a paper reflecting to some
extent the philosophy expressed 30 years earher by
Levitt & Hatnm (1943), questioned whether there was
any physiological advantage in drying back seeds after
treattnetit, or whether this was done simply for con-
venience. He also recognised that the salt solutions
used in seed soak treattnents to 'feed' seeds may simply
have been having an osmotic effect, and he described
the development of the method of seed 'pritning' using
polyethylene glycol solutions, with solute potentials
generally between -1.0 and -1.5 MPa, which bring
seeds to the 'brink' of germination apparently by allow-
ing etnbryo development but not germination. Poly-
ethylene glycols were selected because they appeared to
be true osmotica, free from the complications of ionic or
molecular uptake (as in the case of sugars) or of specific
ion effeets (Uhvits, 1946; Redmann, 1974), but despite
their apparent advantages the method of seed priming
that is now being used for commercial development in
Britain (based on the work of Darby & Salter, 1976) has
been adapted to use salt solutions in place of the mote
viscous polyethylene glycol solutions.
This adaptation is, in fact, very similar to the method
used in the U.S.A. (possibly based on the work of Ells,
1963) and described briefly by Harrington (1969) which
involved soaking seeds in aerated hypertonic solutions of
salts which just, or almost, prevented radicle emergence
from the seed. After the hypertonic soak, seeds were
dried back before sowing, but Salter & Darby (1976)
proposed their method as a means of giving synclironous
germination before sowing the seeds directly in the field
using a fluid carrier.
(3) Effects of imposed treatments
Conclusions regarding seed hydration treatment effects
can differ little today from those reached by Crocker &
Barton (1957) or, indeed, by Kidd & West (1919)
except that it is now recognized that the effect of salts
in the soaking medium can be important. The effect
depends upon the degree of seed hydratioti, tempera-
ture and duration of the treatment, the level of aera-
tion, density of seed mass and whether (possible how)
the seeds are dried back. Treatments used involve soak-
ing seeds in water or on surfaces with a free or restricted
supply of water, or allowitig them to absorb water from
vapour. The treatment may also have involved subse-
quent dehydration, possibly with several cycles of
hydration and dehydration. The treatments and their
effects have been reviewed by Heydecker (1974) and by
Heydecker & Coolbear (1977). Different investigators
have reported favourable, neutral and unfavourable
results. Generally, however, germination or field etner-
gence is more rapid after treatmetit, especially at low
temperature (Bremer, 1928; Chippindale, 1934), the
effect being more pronounced if the seed is not dried
back (Heydecker, 1974). Germination can also be more
uniform or synchronous (Heydecker, 1974; Salter &
Datby, 1976). Germination or field emergenee levels
may also be improved. The increases in field emetgence
of carrot {Daucus carota E.) described by Bremer (1928)
and Currali & Salter (1973) may have resulted sitnply
from the increased rapidity of establishment of treated
seeds in the field, but the results of Chippindale (1934)
for grasses, Eevitt & Hamm (1943) for Taraxacum kok-
saghyz, Hegarty (1970) and Heydecker, Higgins &
Turner (1975) for carrot, Sachs (1977) for watermelon
{Citrullus lutwtus (Thunb.) Matsum. & Nakal.), and
Khan & Knypl (1977) for lettuce (Lactuca sativa L.),
celery {Apium graveolens L.), onion (AUium eepa L.)
and soybean (Glyeine max Merr.) showed clearly that
percentage germination at low temperature ean be im-
proved, whilst Gelmond (1965) for leek (AUium porrum
E.) and Heydeeker et al. (1975) for lettuce demon-
strated that germination at high temperature can also be
inereased. The requirement for alternating temperature

in Dactylis glomerata E. can also be removed by a pre-
hydration treatment (Haight & Grabe, 1972). In addi-
tion to overcoming tetnperature restrictions, seed
soaking treatments have been shown to affect subse-
quent germination under osmotic stress or in saline
conditions. Hajison (1973) repotted faster germination
of pretreated wheat (Tritieum aestivum E.) seeds in a
solution of 0.3 MPa solute potential compared to
untreated seeds, and both rate and level of gennination
of wheat in sahne eonditions have been iticreased by
seed treatments (Chaudhuri & Wiebe, 1968; Idris &
Aslam, 1975). Gray & Steekel (1976) found that a pre-
soak in phosphate buffer solution improved both the
percentage germitiation of lettuce in 0.16 M NH4NO3
solution (c. -0.7 MPa) and seedhng emergence in soil
to which NH4NO3 had been added. Pre-soaking wheat
seeds in water immediately before planting has also
been found to increase seedling emergence from salt-
containing soil (Eyles & Fanning, 1964) and seed germi-
nation on saline media (Chaudhuri & Wiebe, 1968).
Villiers & Edgcumbe (1975) have shown that inter-
mittent periods of imbibition followed by dehydration
can prevent deterioration of stored lettuce seed, and
Basu et al. (1975) have reported (without presenting
data) that a hydration-dehydration treatment extended
the subsequent storage life of a wide range of seeds
under ambient conditions or during accelerated ageing
under different tetnperature and humidity regitnes.
(4) Seed physiology in relation to hydration level
Dry seeds have water potentials of less than 100 MPa
and, except in the case of 'hard' seeds, rapidly imbibe
when placed in good contact with iree water, though the
amount of water taken up depends upon the particular
seed constituents, upon the individual seed and upon
the tetnperature. This first, rapid phase of imbibition is
apparently a purely physical process, occurring equally
in live or dead seeds (Mayer & Poljakoff-Mayber, 1975).
Excess of water may prevent tioimal getmination (see
Section 4.2, below) but if there is adequate aeration the
seeds will germinate. However, full germination does
not require full seed hydration but can occur over a
range of water potentials and hence degrees of seed
hydration, the range being dependent on the particular
species involved (Doneen & MacGillivray, 1943; Hunter
& Eriekson, 1952; McGitinies, 1960; Kauftnann, 1969;
Hegarty, 1976). In addition, a continuous supply of
water to the seed is not essential for getmination to pro-
ceed. MeDonough (1975) has shown for a selection of
grasses, legumes and herbs that itnbibed seeds can be
removed from their souree of water supply before germi-
nation starts, but can still germinate, thus demonstrating
elearly that water held within the seed structure can be
used to pertnit extension growth. Indeed, Stiles (1948)
had shown earlier that different orgatis within the seed
hydrate to different extents, with the embryos in maize
(Zea mays E.) and cotton (Gossypium hirsutum L.)
taking up water to a far greater extent than the other
seed structures, and witli the endosperm in maize and
SEED HYDRATION AND DEHYDRATION 103
the cotyledons in cotton possibly acting as reservoirs of
water for the embryo. The physiology of seeds at hydra-
tion levels leading to normal germination is considered
in Section 4.1.
Seeds may be hydrated via either the liquid or vapour
phases and both have been used to bring seeds to equi-
librium water contents below those that pennit germina-
tion. (Equilibration at very high relative hutnidities
(RH)* is considered separately in Section 4.6). In seed
priming (Heydecker, 1974), an osmotic solution is used
to control water uptake whereas in seed 'hardening' or
'advancing' (Austin, Eongden & Hutcliinson, 1969), a
known quantity of water is added to the dry seeds to
bring them to the required water content. In both these
treatments the seeds are sufficiently liydrated to permit
seed activation without deterioration, but if seeds are
hydrated to ratlier lower moisture contents ('damp'
seeds) they deteriorate rapidly. Seed storage deteriora-
tion work often uses vapour equilibration to eontrol the
level of seed hydration. The physiology of activated
seeds is considered in Section 4.4. and that of 'damp'
seeds in Section 4.5.
The extent of seed hydration at a given water poten-
tial is dependent upon the character of the particular
seeds, varying with the species and seed constituents
(Owen, 1956; Crocker & Barton, 1957; Roberts &
Roberts, 1972) as well as with individuals within a seed
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
liigh enough (over 43%t), non-dormant seeds will germi-
nate; in the range from 22 to 43%, heating due to micro-
organisms will occur if oxygen is present, resulting in
rapid death of the seeds; in the range from about 11%
(oily seeds) or 15% (starchy seeds) to about 22%,
storage fungi grow actively and destroy the seed embtyo.
These last values represent moisture contents in equi-
librium at 20C to 25C with RH's of 65-70%) at the
drier end, and 85-90% at the wetter end of the range
(Christetisen, 1973). Thus this view essentially proposes
three effective ranges: fully itnbibed seeds which will
germinate; damp seeds that deteriorate in association
with microorganisms; and dry seeds. This simple view
fails to represent the data adequately, and instead
Hegarty (1977b) has proposed five ranges of seed hydra-
tion which take into accoutit in addition seed activation
without gennination, and seed damage in very dry con-
ditions, the latter being discussed in Section 4.3.
As an addendutn to the introduction to tliis Section
it is wotth etnphasizing the nature of the relationship
between RH, water potential and seed water content.
At 15C, a water potetitial of -0.15 MPa is equivalent
* The preferred units for describing the moisture eontetit of
air are g m"' rather than % RH. The former is, however, tem-
perature dependent, and as temperature data have not been
included in all the papers in which RH figures are given, conver-
sion of % RH to g m '' is not possible in all cases. For consis-
tency, % RH has been used throughout.
t Seed tnoisture eontents are expressed on a dry weight basis
and data from papers have been adjusted to this base where this
has been necessary.
104 T. W. HEGARTY
to 99.9% RH; -1.5 MPa to 98.9% RH; -15.0 MPa to
89.5% RH; and -150 MPa to 33% RH (Slatyer, 1967).
In very approximate terms a seed may contain half as
much water again at 0 MPa as at 1.5 MPa (Hegarty,
1977a) so that at equilibrium the seed would be two-
thirds hydrated in an atmosphere of 99% RH, the
remaining one-third of potential water uptake being
associated with a change of only 1% RH (if hydration
by vapour and liquid can be equated).
(4.1) Seeds hydrated sufficiently to permit germination
During the germination process, water uptake occurs
essentially in three phases, with a more or less distinct
static or lag phase separating two phases of rapid water
uptake, the first associated with initial imbibition and
the second with radicle growth. The extent of the lag
phase is dependent upon the species, individual seed and
temperature (Berrie & Drennan, 1971; Hegarty, 1977a),
but although water uptake may be negHgible during
this phase, the seed is not inactive.
According to Brown (1972), it used to be commonly
accepted that the stage up to the rupturing of the seed
coat by the radicle was more or less passive, with events
determined by the physical properties of the system.
Instead we should now recognize the pre-germination
phase as including: an activation of enzymes that are
present in the dry seed; a change in the structure of the
cytoplasm that sustains metabolic activity; a synthesis
of enzymes both before and after active growth has
begun; and a change in the metabolic pattern as a
result of a change in the quantitative relations between
enzymes in the growing system.
This pre-germination phase has itself been subdivided
into phases by a number of different authors. Evenari
(1961) describes four phases (imbibition; activation;
mitosis; and protrusion) whilst Berlyn (1972) also sug-
gests four phases, though with a slight difference in
emphasis (imbibition; hydration and activation; cell
division and extension; and protrusion of the embryo
from the seed). Perhaps more closely correlating with
the pattern of water uptake and respiration in seeds is
the less rigid description of the germination process by
Ching (1972; 1973) in terms of three distinct but over-
lapping phases. The first (A) involves the hydration and
reactivation of existing systems when there is likely to
be a start to basal metabolism, an increase in ATP con-
tent for various synthetic activities following imbibition,
and the provision of substrate for respiration and protein
synthesis. The second phase (B) involves synthesis of
enzymes and organelles for catabolism of reserves and
occurs mostly in the storage organs, with the activity
or quantity of the protein-synthesizing machinery, mito-
chondria, glyoxysomes, enzymes, coenzymes, cofactors,
substrates and other factors increasing during and after
imbibition. The third phase (C) involves synthesis of
new cellular components and is associated with radicle
emergence from the seed, using substrate transported
from the storage organs. Both phases B and C are likely
to be affected by environmental influences, possibly
through effects on reactivation and, in the case of phase
C, on catabolism.
The rapidity of reactivation of pre-existing systems
has been emphasised by the work of Marcus, Feeley &
Volcani (1966) who found in wheat embryo that the
rise in protein-synthesising capacity lagged only 10
minutes beliind water uptake at room temperature. The
independence and interdependence of the relationsliips
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
growth of decotyledonized radish {Raphanus sativus L.)
embryos, and by Sutcliffe & Bryant (1977) in their
description of seed germination in the pea {Pisum
salivum L.). From these descriptions it is clear that
developments in phases B and C are dependent on pro-
cesses occurring in their preceding phases, and in radish
it appeared that even witliin a single phase (B), progress
was dependent upon processes occurring in an early
period of that phase. However, respiratory activity in
phase B was not affected in conditions wliich inhibited
respiration in phase C.
For continuing radicle growth there must be concur-
ring cell division and cell extension, although their onset
may not be simultaneous (Berlyn, 1972). Rogan &
Simon (1975), using their own data and the results of
other workers, describe an initial slow increase in root
length in broad bean {Vicia faba L.), barley {Hordeum
vulgare L.), pea and French bean {Phaseolus vulgaris
L.) which is followed by a phase of rapid growth co-
inciding approximately with the onset of mitosis.
According to Sutcliffe & Bryant (1977), increase in the
volume of the radicle in pea causes it to emerge from the
testa before the start of the phase of rapid growth (tliis
latter involving a large increase in DNA and RNA con-
tents of the axis accompanied by rapid cell extension).
This coincidence of cell division and rapid cell elonga-
tion has also been found in oat {Avena sativa L.) and
tomato {Lycopersicon esculentum Mill.) (Berrie &
Drennan, 1971) and in lettuce at 26C (Evenari et al.,
1957). However, Haber & Luippold (1960a), whilst con-
firming the results for lettuce at 26C, were able to show
that cell division and cell elongation were not necessarily
synchronous, but could be delayed with respect to each
other by low temperature (10C delaying mitosis) or
by osmotic stress (0.4-0.5 M mannitol, 1.0 to 1.3
MPa, delaying elongation). Under extreme circumstances
(heavily irradiated seeds) plantlets of some species
capable of CO2 fixation and dry weight increase can
grow, even thougli mitosis has been completely sup-
pressed (Berlyn, 1972). TIius radicle protrusion from the
seed may be due to cell elongation in combination with
cell division or to cell elongation alone wliich can result
in 'false' germination, a phenomenon that is considered
in Section 8,below.
Respiratory patterns during the germination process
are complex and have been discussed by Roberts (1969,
1973a) and, briefly, by Mayer & Shain (1974). In dry
seeds mitochondria have been found to be poorly dif-
ferentiated and lacking in malate dehydrogenase, but as

imbibition proceeds they become rich in protein and
Upid, and biologically active (Nawa & Asahi, 1971). A
shift in respiratory pattern 4-8 h after imbibition has
been found in soybean, oat, lettuce, corn, tomato and
pea, from predominantly alternate respiration to a
cyanide-sensitive respiration (Yentur & Leopold, 1976),
and Howell (1961) reported alterations in metabolic
characteristics of soybean cotyledon mitochondria
during the course of the germination process. Toole et
al. (1956) have suggested that in a physiological sense,
the start of germination depends upon coupHng respira-
tion to growth. However, whilst radicle growth may be
linked with increased respiration, respiration can pro-
ceed at a substantial rate in the absence of growth, i.e.
in seeds that for one reason or another cannot germinate
(Ching & Foote, 1961; Chen & Varner, 1970; Hegarty,
1977a) and long before growth begins, energy is coupled
to endergonic metabolic processes (Haber & Luippold,
1960b). However, the extent to which this respiration is
associated with embryo development or repair mechan-
isms, and the extent to which respiration is uncoupled,
'awaiting' growth, is not clear.
(4.2) Soaked seeds
Seeds immersed in water imbibe, may lose a wide range
of solutes and, in time, may germinate or suffer damage.
Although many seeds will not germinate under water,
some certainly can, in a very restricted supply of oxygen
(Carr, 1961). In an extensive investigation, Morinaga
(1926) placed seeds in Erlenmeyer flasks filled with
distilled or boiled water. He found that germination of
18 species was virtually the same under water as on filter
paper, 25 species showed reduced germination, and 34
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
of reserve material in the seeds. An atmosphere of oxy-
gen rather than air in the flasks gave liigher levels of
germination.
According to Carr, when seeds arc soaked in water
they go through a period of anaerobiosis which, if too
protracted, can cause injury, resulting in inability to
germinate or in poor germination and ready decay. The
state of tlie seed and the environmental conditions seem
to be important. For instance, dormant seeds appear
not to be damaged excessively by storage in water;
increased injury in some species at 10 or 30C compared
to 20C is associated with greater losses of solutes from
the seeds; and many kinds of seeds are injured by soak-
ing in oxygenated water but may be protected by the
presence of hydrogen peroxide or, more readily, by
carbon dioxide. In Frencii bean. Barton & McNab
(1956) found that oxygenation caused excessive water
uptake in the seeds, and after extended (24 h) soaking
in the presence of oxygen there was also excessive
leaching from the seeds.
Crawford (1977) has suggested that seeds resistant to
soaking injury regulate glycolysis so that there is mini-
mal production of ethanol, and that the toxic effects of
SEED HYDRATION AND DEHYDRATION 105
ethanol are due to a deleterious 'fluidizing' of the cell
membranes. Certainly, the damage associated with the
leaching of solutes at higli and low temperatures or in
the presence of excess oxygen, described above, would
indicate a general role of membrane involvement in
soaking injury.
(4.3) Very dry seeds
Over-desiccation can damage seeds (Harrington, 1972;
Ching, 1973; Woodstock, Simkin & Schroeder, 1976).
Harrington (1972) suggests that water in seeds can be
described in terms of three categories: unbound, mobile,
free, capillary or solution water; multilayer, chemi-
sorbed, hydrogen-bonded, intermediate water; and
monolayer, localized, polar-site, bound water. As seeds
are dried, first the free, then the intermediate water is
lost, and below 25% RH the monolayer water is progres-
sively lost from the macromolecules, removing a layer
that is protective against oxidative processes. The most
serious oxidation is the autoxidation of lipids which can
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
antioxidants but these are used up during storage as
oxidation occurs and no more are produced in very dry
seeds.
Whilst it might be expected that withdrawal of mono-
layer water would itself affect macromolecular structure
and function, the free radicals produced can damage
cellular membranes and can react destructively with
macromolecules, destroying protein, DNA, and cell
reproductive capacity (Harrington, 1972; Harman &
Mattick, 1976).
(4.4) Seeds sufficiently imbibed to be activated but
unable to germittate (incompletely imbibed seeds)
Data on the pliysiology of incompletely imbibed, non-
dormant seeds are sparse and scattered, and the picture
of what is happening in such seeds must to a large extent
be inferred from work such as that of Heydeeker et al.
(1975) and Salter & Darby (1976) that seeds can be
brought to the 'brink' of germination if under the ap-
propriate osmotic control. The very rapid germination
on transfer to water, and the improved uniformity of
germination imply that the germination process in these
seeds has proceeded via one route or another until a
particular block is reached (i.e. phases A and B are
nearly if not altogether completed, thougli perhaps at a
slower rate than in water), and that the seeds within a
single population reach tliis block at different rates but
germinate relatively uniformly once the block is released.
The data presented by Hagarty (1977a) on the
respiration of calabrese {Brassiea oleraeea var italiea
Plenck.) and carrot seeds when held in solutions of dif-
ferent solute potentials showed that in the lag phase of
the germination process (phase B) the rate of oxygen
consumption was dependent on the degree of seed hy-
dration, but even at hydration levels which just pre-
106 T. W. HEGARTY
vented germinafion, oxygen uptake rates were relatively
higli. Malnassy (1971) found that the addition of cyclo-
heximide (an inhibitor of protein synfhesis) to the seed
soak medium (ati aerated solution of NaCl at c. 1.0
MPa) partly or completely negated the beneficial effect
of the seed soak on subsequent germination of pepper
{Capsicum annum L.), celery and totnato seeds. He also
found isoenzyme differences in treated and untreated
seeds, and quoted the work of Koehier (1967) showing
that levels of RNA increased 80%, and peroxidase levels
increased eightfold in tomafo seeds during the treat-
ment. High nucleic acid levels in 'primed' tomato seeds
were also reported by Coolbear & Heydecker (1975).
Using 'pdmed' seeds of lettuce, Klian et al. (1978)
have shown that, compared to untreated seeds, there are
iticreases in the rates of ^H-uridine iticorporation into
RNA, polyribosome incorporation and '''C-leucine
incorporation into profeins; iticreased activities of sotne
enzytnes thougli not of others; and qualitative and
quantitative changes in soluble proteins and isoenzymes
of esterase, 3-phosphoglyceraldehyde dehydrogenase,
and phosphatases. In addition, a 2-week osmotic treat-
ment at ]5C led to a complete disappearance of ab-
scisic acid.
Subsequent reliydration after dehydration has re-
sulted in higher respiration rates in treated compared to
untreated seeds (Koehler, 1967; Woodstock, 1969;
Malnassy, 1971), suggesting that substrates may be
accutnulated, or systetn activity may be enhanced in
seeds that are just prevented from gennination.
In addition, cell division has been shown to occur in
carrot embryos at seed hydration levels that almost
certainly would have prevented gennination (Austin ct
al., 1969) and in embryos of lettuce seeds held in
mannitol solution (c. 1.3 MPa) which virtually pre-
vented germinafion (Haber & Luippold, 1960a), wliilst
cells in a metabolically active state were detected in
radicles of Nemnphila mcnziesii Aggr. held in mannito]
solution of c. -2.0 MPa (Cruden, 1974).
Thus considerable activity is possible in partially
hydrated seeds, including, apparently, many or all of the
processes of phases A and B of fhe germination process,
and possibly even the operation of systems permitting
cell division, but not cell elongation co-ordinafed with
cell division. It has not yet been shown, however,
whether the physiology of incompletely imbibed seeds
is essentially similar to that of fully imbibed seeds up fo
a particular point and then diverges, or whether the
patterns of development in these fwo situations are quite
distinct.
(4.5) 'Dry'and 'damp'seeds
At room temperature 'dry' seeds of tnany species can be
stored satisfactorily, but even af the optimum moisture
content for storage, seeds will deteriorate wifh age, the
rate of deterioration (Roberts, 1972) and metabolistn
(Bailey, 1940; Edwards, 1976) increasing as fhe seed
moisture content is raised. Thus the optimutn moisture
contenf for dry storage represents a balance between
minimizing the deterioration resulting at the moisture
level of 'damp' seeds and that caused by excessive desic-
cation. Bass (1975) suggests equilibrium with 30% RH
as being best for many seeds, whereas Harrington (1972)
suggests equilibrium wifh 20-25% RH as the opfimutn.
Changes that occur during storage and are associated
with deterioration lead to delayed gennination, reduced
seedling growth rates, decreased tolerance of adverse
germination conditions atid loss of germinability (Abdul-
Baki & Anderson, 1972; Delouche & Baskin, 1973).
However, the underlying causes of these chatiges are tiot
clear, tiot so much because no cytological and metabolic
changes have been detected, but because so tnany have
been recorded (Roberts, 1973b). Roberts described
sotne of the tnore consistent observed changes in seeds
which include: cytological damage, including datnage to
the chromosomes and ribosotnes sytnptomatic of a
general degradation of nucleic acids, damage to other
organelles symptomatic of general degradation of hpo-
protein tnetnbranes, atid datnage fo the cell tnetnbratie
itself; alterations in the activity of enzytnes, particularly
those concerned with redox activity which decrease in
activity, and sotne hydrolytic enzymes which increase in
activity; various changes in cell consfituenfs includitig
profeins, fatty acids, reducing and tion-reducing sugars
and growth inhibitors; and a decreased ability to incor-
porate isofopically labelled sugars and amino acids.
Some of these changes ate accelerated by micro-
organisms, but Roberts considers that they can also
occur in their absence and in any case are compatible
with the general increase in hydrolyfic activity.
Accelerated ageing tests set out specifically to ac-
celerate the processes of nonnal ageing by exposing the
seeds to high RH and temperature for a specified period
of titne. In general, the tnost usual conditions are 100%
RH at 40-45C with exposure titnes of from 2-8 days,
but in some cases a less severe regime of 30C and 75%
RH for periods of 6-24 weeks is used (Delouche &
Baskin, 1973). Loss of vigour precedes loss of viability,
but fhe test itself is concerned only with nortnal gertni-
nation after ageing, with high vigour seed lots showing
less reduction iti germination than low vigour lots.
There is much evidence that deferiorafion in 'datnp'
seeds is a physiological process, but Christensen (1973)
has argued that there is sufficient evidence fo itnplicate
fungal activity as a primary cause of seed deterioration,
and he is careful to distinguish between the effects of
storage fungi, active in RHs of from 70-90%, from those
of field fungi, active in excess of 95% RH. However,
Harrison & Perry (1976) have presented convincing
evidence for fhe role of physiological deterioration of
barley held in damp store. Deferioration increased with
iticreasing seed moisture content, and at most moisture
contents the start of decHne in vigour and viability
clearly preceded any substantial invasion by fungi.
Deterioratioti in this case was measured in terms of
reduced membrane integrity, atid both reduced respira-
tion and ability to synthesise a-atnylase during sub-
sequent germination.
Loss of membrane integrity seems to be intimately

involved in the deterioration process (Koostra, 1973)
and is probably a primary step in the sequence of events
during detetioratioti (Delouche & Baskin, 1973).
(4.6) Seed responses at very high relative humidities
If water potential predominantly controls water uptake
in seeds, one wotild expect sitnilar results of seed treat-
ment whether carried out in contact with a water-
supplying substrate or in controlled atmospheres. The
exception to this would be if physical hydration were
different for some reason in the two situations, as has
been suggested by Shull (1916), or if the rate of supply
of water were importatit. Much of the work on seed
'pritning' has involved the use of solutions in the range
1.0 to -2.0 MPa and, as has been pointed out earlier,
an osmotic solution of -1.5 MPa at 15C is in equi-
libriutn with a RH of c. 99%. Results of investigations
concerning the effects of saturated or nearly saturated
atmospheres on seeds are widely contradictory, with
reports of seed germinatioti in some cases, activation
without germination in others, and deterioration to a
greater or lesser extent in a thitd group.
Equilibrium tiioisture contents attained at 100%
RH have been quoted for a range of crops (mainly
cereals) by, for instance, Coleman & Fellows (1925)
and by James (1967) witli values ranging from approxi-
tnately 27% in flax {Einum usitatissimum L.) to 37% in
barley, the values quoted in the two papets agreeing
closely. Coleman & Fellows used a fiow of saturated air
rather thati a static systetn. No tnention was ttiade in
either case of fungal growth or germitiation occurritig.
Deterioration of seeds in saturated atmospheres has
been described by Robertson, Lute & Gardner (1939),
Brewer & Butt (1950) and Griffin (1966) whereas other
reports describe germination (Black, 1959; Owen, 1952)
and improvemetits in subsequent seed gertnitiation
(Levitt & llatnm, 1943; Wilson, 1973; Heydecker,
1974). Accelerated ageing treatment tnay involve high
hutnidity storage and results in seed deterioration
(Delouche & Baskin, 1973). Fitially, Villiers (1974)
compared lettuce seeds hydrated in water vapour for 2
weeks befote sealed storage, with seeds kept fully im-
bibed but thermodormant and found that the former
showed deterioration whereas the latter did tiot. He
noted, however, that the vapour-equilibrated seeds had
attained a moisture content of only 14%.
Apart from the possible different responses of indi-
vidual species or seed lots to being held in saturated
water vapour, four explanations of these differences
seem possible. First, it is extremely difficult to main-
tain temperature conditions so constant that the atmo-
sphere remaitis saturated yet no condensation of water
on the seeds occurs. Machalica (1926) has reported that
germination is greater in a moist atmosphere with fluc-
tuating compared with constant tetnperature, and
fiuctuating temperature has also been shown to increase
seedling emergence from dry soil, probably through an
effect on the water vapour relatiotis of germitiation
(Hegarty, 1975). Second, Christensen (1973) has pointed
SEED HYDRATION AND DEHYDRATION 107
out that absolute uniformity in moisture content is
almost itnpossible to attain even in a stnall satnple of
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
differences in oxygen or carbon dioxide concentrations,
especially around seeds held in confined atmospheres,
could weD have influenced the results. Fitially, the
physiological response of the seed tnay be determined
by the rate of increase of hydration and by the absolute
level of hydration when in equilibrium with saturated
water vapour. If the rate of increase of hydratioti is slow,
the seed may go througli a protracted peiiod at hydra-
tion levels that favour deterioration rather than activa-
tion and repair, and although a higher tetnperature
would favour more rapid hydration, it would dso favour
tnore rapid deterioration. In addition, seeds if truly
equilibrated with saturated vapour may well be sub-
stantially less hydrated than if itnbibed in contact witli
water in the liquid phase, especially if there are parts of
the true seed or outer coverings that can retain quanti-
tities of 'free' water which can be utilized by tlie seed
for cell elongation during germination.
Thus discrepancies reported on the effects of high
RH on seeds could well relate to the absolute level and
rate of hydration of the seeds, to the degree of tem-
perature control over the environtnent in which the
seeds are held, and to the levels of carbon dioxide or
oxygeti maintained in that environment. However, it
does seem possible that, in vapour, under carefully
controlled temperature conditions, seed activation and
germination can occur over a range of water potentials
(Owen, 1952).
(4.7) Conclusions
Ouite distinct physiological processes occur at different
seed moisture contents, although they are not well
understood and are at best recognized through their
effects on seed constitution or on perfortnance during
germination. In 'very dry' seeds, datnage tnay be the
result of excessive removal of water frotn the seed; in
'dry' seeds, deterioration occurs but its rate is tninimal;
in 'datnp' seeds, both metabolism and deterioration ap-
parently increase with water content; at higher water
potentials there must be a transition point when repair
reactions are activated or co-orditiated, or metabolism
becomes linked with seed activation, so that activatioti
and repair rather than deterioration are dominant.
Substatitial etnbryo developtnent can occur in this state
of 'incomplete' imbibition. Only at higlier water poten-
tials does cell extension occur with cell division to per-
tnit continuitig radicle growth. Even then, full itnbibition
is not essential, there being a range of water potentials
over which germination will occur.
(5) Differences between individual seeds
There now seems to be genetal agreement with the view
expressed by Hunter & Erickson (1952) that seeds must
108 T. W. HEGARTY
reach a critical level of hydration to permit germination.
It does not, however, seetn to be equally generally ap-
preciated that individual seeds eveti within a seed lot
can hydrate to quite different levels before gennination
commences.
In a detailed study of water uptake in calabrese seeds
placed in contact with polyethylene glycol solution of
-0.75 MPa at 10C (Hegarty, utipublished) the indi-
vidual seeds were weighed at frequent intervals for 35
days or utitil germitiation occurred. All seeds showed a
lag phase in hydration level after initial imbibition and
before the start of germination, and of 50 seeds used 32
germinated between 11 and 35 days after the start of
imbibition and a further 15 seeds gertninated normally
on transfer to water after 35 days. Of the 32 seeds that
germinated at -0.75 MPa the mean water uptake just
before germination relative to the original seed weight
was 54%, with a range of from 44 to 66%, whereas the
tnean water uptake of the other 15 seeds after 35 days
was 62% (significantly higher than 54%) with a range
of from 50 to 83%. Thus although the ranges of seed
water uptake overlapped, the seeds that did not germi-
nate at 0.75 MPa bars were in general more hydrated,
not less, than those that did, and would only germinate
in an environment of higher water potential. Owen
(1952) also found that non-germinated seeds of wheat
had higher moisture contetits than those that had
gertninated.
A different facet of this situation is expressed in the
germination responses of seeds to moisture stress in
which, for a particular lot of seeds, germination is de-
creased progressively over a range of decreasing water
potentials until no germination at all can occur. Relevant
to this is the statement of Crocker & Barton (1957)
that low vigour seeds may need tnore water to germinate
compared to high vigour seeds, and it is also relevant
that seed deterioration or loss of vigour is claimed to
lead to a diminution in the range of conditions in which
seeds can germinate (Abdul-Baki & Anderson, 1972).
Thus it is not the level of hydration per se that governs
whether or not a seed will germinate.
The work of Stiles (1948) on the distribution of
water within seeds has already been mentioned and it
would clearly be of considerable interest to know how
reduced water potential affects the hydration levels of
the various seed structures, especially in individual
seeds, and how it might also affect the inter-relation-
ships between the embryo and its surrounding or en-
closed reserve materials.
(6) The effects and timing of dehydration
Pre-germination hydration treatments can result in tnore
rapid subsequent germination than in untreated seeds,
although this may be partly due to more rapid imbibi-
tion resulting from a change in the physical structure of
the seed or seed coat (Kidd & West, 1919; Heydecker
et al., 1975; Heydecker & Coolbear, 1977). There is,
however, increasing evidence that the effect of physio-
logical development in the seed need not be lost on de-
hydration, althougli the beneficial effects tnay be
counteracted by damage, either gross or more subtle,
or by other responses on dehydration.
In peas, during the initial stage of rapid hydration,
the resistance of the seed to dehydration remains lugh,
but subsequent uptake of water to the hydration level at
germination causes a rapid decrease in dehydration
resistance, apparently associated with the accumulation
of water in cytoplastnic vacuoles (McNair, 1966, quoted
by Sutchffe & Bryant, 1977). Dehydration of oat seeds
following imbibition and prior to the first signs of visible
germination has no effect on level or rate of gertnination
on subsequent rehydration, suggesting that the titne
required for cotnpletion of germination by a population
of seeds depends upon the total time they are hydrated,
irrespective of whether or not this is interrupted by a
dry period (Akalehiywot & Bewley, 1977). However,
Berrie & Dretman (1971) considered that the effects of
hydration before dehydration were only cumulative in
oat seeds if the hydration period was longer than a
certain minitnum duration.
Sen & Osborne (1974) found that getmination in
low-viability rye {Secale cereale L.) etnbryos was en-
hanced by a suitable hydratioti-dehydration treatment of
the etnbryo, and that there was a balance between the
advancement of synthetic processes due to hydration
and datnage to the earUest activated cells incurred
during subsequent dehydration, where datnage was indi-
cated by the loss of ability to incorporate leucine
(protein synthesis) or thymidine (DNA replication).
The hydration pretreattnent initially enhanced the
ability of the embryo to synthesise protein and RNA
compared to the untreated controls, and the nortnal 9 h
lag before the onset of DNA rephcation could be abol-
ished. Thus at least some of the changes occurring in the
initial stages of germination were stable to dehydration
and subsequent storage. Support for this suggestion
comes frotn the earlier work of Marcus ct al. (1966) who
showed that desiccation did not destroy ribosomal ac-
tivity in wheat embryos, and of Chen, Sarid & Katchal-
ski (1968) who also investigated hydration-dehydration
treatments in wheat embryos. They found that rehydra-
tion for 24 h led to the transcription of new and active
mRNA in embryos that had been previously hydrated
for 24 h but in those previously hydrated for 72 h only
trace atnounts of complementary RNA were transcribed,
and the transcribed RNA differed from the mRNA
fonned in normally germinating embryos. It was sug-
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
during the early stages of gennination. Thus it is pos-
sible that datnage on dehydration is associated with
DNA rephcation and RNA transcription.
This more subtle datnage to the etnbryo should per-
haps be distinguished from gross damage incurred when
the growing embryo is dehydrated, although the two
may be related. The Russian work on seed 'hardening'
(May et ah, 1962) suggested that the latest stage at
which seeds should be dried back was at the time of

radicle emergence frotn the seed (i.e. as cell extension
growth comtTiences, and possibly also coinciding 'with
the start of cell division, c.f. Bertie & Drennan, 1971)
and Hubac (1962) has said that the ability of seeds to
survive dehydration is lost as hydration of the vacuoles
of the embryo eommenees. Ho'wever, Carcellar & Sori-
ano (1972) dried back wheat seeds just after germination
without deleterious effeets on subsequent germination,
and even if this is done repeatedly, viability may be re-
tained in some seeds over several cycles of hydration
and dehydration, with cereals being more resistant than
peas (Widstoe, 1916). Seedlings at an even more ad-
vanced stage of germination can be dehydrated without
causing death, although growth may be resumed from
secondary initials (Milthorpe, 1950; Hegarty, 1977a).
It is interesting that isolated embryos of low-viability
rye when at too advanced a stage before dehydration
also initiated growth from secondary initials (Sen &
Osborne, 1974).
In oats, Berrie & Drennan (1971) found that protease
activity was higher in treated seeds on subsequent im-
bibition than untreated seeds, that this response was
stable in stored seeds and that the effect of successive
hydration-dehydration cycles was cumulative only if the
prior imbibitions were of substantial duration. In crested
wheat grass (Agropyron desertotttm (Fisch. ex Link)
Schult), Wilson (1971, 1973) found that incubation of
seeds in vapour over solutions with water potentials
down to -4.0 MPa, or seeds placed in soil and recovered
at various times during the gertnination process, showed
an increased subsequent rate of germination at low tetn-
perature, with the increase in a-amylase activity being a
good indicator of the hastening effect of the treatment.
In a series of eight cycles of hydration-dehydration, a-
amylase activity was successively accutnulated without
loss on dehydration. In contrast, seeds sown on the soil
surface showed increase ATP levels after rainfall but
these levels fell again under severe tnoisture stress
(Nelson, Wilson & Goebel, 1970).
Futther evidence for either accumulation of sub-
strates or enhancement of activity of enzymes or systems
in seeds prevented from germination by osmotic stress,
comes from the subsequent inereases in respiration rate
after dehydration and imbibition compared to non-
treated seeds. This has been reported for pepper (Wood-
stock, 1969; Malnassy, 1971), tomato (Woodstock,
1969; Koehler, 1967), and Dactylis glomerata (Haight &
Grabe, 1972). This effect might resemble the 'stored
growth effect' described by Ray (1961) in which oat
coleoptile growth is reduced by transfer to osmotica but
then surges forward itiitially at a much greater rate on
transfer to water. This is apparently not simply a physi-
cal effect associated with osmotic properties because the
stored growth effect can be inhibited by KCN.
One further response resulting in the negating of
beneficial effects of seed 'priming' of celery has been
described by Gibbins & Heydecker (1977) in which a
germination inhibitor apparently accumulates in the
seed on dehydration, causing a form of secondary
dormancy.
SEED HYDRATION AND DEHYDRATION 109
The effects of dehydration on germination response
described above seem to fit very well the responses of
over 40 grass, weed or woody species of Utah range
plants subjected to alternate wetting and drying cycles
(Griswold, 1936). Griswold concluded that prior to ger-
mination, interruption of the moisture supply may
arrest the development or growth of the embryo which
precedes germination until sufficient moisture is sup-
plied again, whereas dehydration may alter the proper-
ties of the seed coat, may produce secondary dormancy
or, if at a critical stage, may injure the embryo beyond
the point of recovery.
One final effect that may be of relevance is the
phenomenon of vernalization which is induced in seeds
held fully or partly imbibed in the cold (Napp-Zinn,
1961). The phenomenon is apparently reversible,
althougli the loss of the effect when it does occur may
be due to too high a temperature being maintained
during dehydration rather than to the process of de-
hydration itself (Schwabe, 1971). There seems to be
no physiological advantage in using the Russian method
vvhicli prevents germination during the treatment, tliis
being considered to be simply more convenient for
subsequent handling of the seed (Purvis, 1961).
Thus some at least and possibly many of the physio-
logical developments associated with the preparation of
the seed for gerrnination appear to be stable to de-
hydration if they are not counteracted either by cell or
organ damage in the embryo (which may coincide with
the initiation of cell elongation or cell division) or by
some form of secondary dormancy. It seems possible
that an important difference between pretreatment in a
restricted quantity and in a free supply of water eould
be the extent to which substrates can accumulate or
system activity can be enhanced in the seed for use
during subsequent germination.
Also relevant in this Section is the contrast made by
Thomas (1972) between the process of dehydration in
maturing seeds and rehydration after maturation. Some
of the marked changes in cell organisation wliich occur
as the seed matures are direct expressions of changes in
the state of cell membranes atid may be related to bio-
chemical changes in dehydrating seeds. The maturing
seed is subjected to water stress probably severe enough
to contribute to marked changes in respiratory activity
and enzyme synthesis, and Thomas accepts the sug-
gestion of Klein & PoUock (1968) that ehanges in the
physiological activity of the tnaturing seed are adapta-
tions which render the cells resistant to desiccation. In
addition, he poses the question whether water stress
might alter growth regulator levels in maturing seeds (as
it does in leaves), with an implied control over metabol-
ism. He also points out that the proteins synthesized by
maturing seeds are very different to those made by
germirtating seeds, and presumably the mRNA's must be
different in tlie two cases. It follows that at some stage
during seed development there must be destruction of
the old messetiger and a synthesis (and possibly storage)
of the new, and it is possible that desiccation, by
directly affecting cell nuclear volume and the osmotic
110 T. W. HEGARTY
and ionic environment of the chromatin, causes the
ordered openitig up or closing down of genetic areas.
If Thomas's hypothesis is correct, then, by analogy
with the maturing seed, it is possible first that at sotne
stage during the germination process or during the seed
priming treattnent the etnbryo may lose its adaptation
to desiccation and hence suffer damage if subjected to
dehydration, and second, that desiccation during the
gertnination process may itself cause a change in genetic
response to subsequent rehydration.
One possible site of damage during desiccation is the
cell metnbrane and, in his review on freezitig injury in
cells, Merytnan (1974) argues persuasively that solute
concentration damage may not be the result of de-
naturation by the concentration of any specific solute,
but appears to be related to the osmotic reduction of
cell volume, with the possibility that the cell, and par-
ticularly the tnembrane, is damaged mechanically if a
cell decreases in size below a tninimum critical volume.
He suggests that there is a mechanical resistance to
volume change in cells associated with the metnbrane
and, as a result, the internal hydrostatic pressure of the
cell matrix contained within the plasmaletnma can fall
below attnospheric pressure, thus creating a pressure
gradient across the membrane. This, in turn, can lead
to an abrupt change in permeability of the metnbrane
(the site of pritnary damage), permitting an influx of
extracellular solutes and loss of stability of the metn-
brane macromolecules, thus pertnitting further
degradative events leading to irreversible change.
Although this hypothesis is proposed to account for
the effects of freezing injury in cells, with the etnphasis
being on animal cells, it clearly has itnpHcations for the
process of dehydration in seeds after hydration, and, if
cell shrinkage were differentially affected, could offer
an explanation for differences in response due to dif-
ferent rates of drying.
(7) Analogous situations
(7.1) Salt marsh species
The closest analogy to the seed 'priming' treatment
occurs with seeds released frotn salt tnarsh plants into
sea water (Chapman, 1960; Boorman, 1968). Seeds of
the tnajority of halophytes, although apparently resis-
tant to the damaging effects of NaCl, germinate best in
fresh water conditions. If first held in sea water, they
germinate very rapidly on transfer to fresh water. Boor-
man considered that the early stages of the germination
process could take place in salt water and only the final
stages preceding germination were inhibited.
(7.2) Arid environments
Successive cycles of wetting and drying can occur in
arid environments and are likely to permit developtnent
of the embryo, resulting in more rapid germination
either when substantial rainfall occurs, or at low tem-
peratures in the spring. Rapidity of establishment in
either set of circumstances may give a survival advantage
to seedlings (Wilson, 1971; Wilson, Wotidercheck &
Goebel, 1974). In less extreme environtnents, tnoisture
stress can delay seed germination in the soil and can
result in a double fiush of seedlings after subsequent
rainfall. A laboratory experiment with calabrese and
carrot seeds suggested that pre-etnergence losses in the
particular conditions used were due to seedling dehydra-
tion rather than to datnage of the seeds (Hegarty,
1977b), although the precise degree of seed hydration
may be very important, especially with respect to the
interaction of the seed with seed or soil pathogens
(Wallace, 1960).
{1.3) Dormartcy
Several types of dortnancy yield situations in which
there is active seed metabolism yet no gertnination. In
some species, after-ripenitig must take place in itnbibed
seeds before specific dormancy-breaking treatments
leading to germitiation are started. In such seeds, ana-
tomical and morphological changes tnay occur (Mayer &
Poljakoff-Mayber, 1975). Detailed studies in ash {Fraxi-
tws excelsior L.) have revealed tliat althougli no cell
division occurs during the after-ripening phase, there are
extensive food cotiversions, wall development and the
appearance and proliferation of cellular organelles
duritig this phase of deep dormancy (Villiers, 1971).
In dormant seeds sliortly after imbibition, rates of
respiration as high as or approaching those of non-
dormant seeds have been found in wheat (Miyamoto,
Tolbert & Everson, 1961 ;Ching& Foote, 1961), in wild
oat {Avena fatua L.) (Chen & Varner, 1970) and in
lettuce (Foutitain & Bewley, 1976), suggesting that
seeds are active at that time though they cannot gertni-
nate. In barley and rice {Oryza sativa L.), initial rates of
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,
1968).
Seeds can survive, imbibed, in the soil for long
periods (Crocker, 1916) yet respiration rates of such
seeds can be appreciable (Barton, 1945), adding support
to the theory that seeds in soil tnay eventually die due
to exhaustion of food reserves (Crocker, 1916). The
rate of respiration found for non-dormant calabrese
seeds held under moisture stress in polyethylene glycol
solution seemed to be cotnpatible with survival for a
long period before the seeds' reserves would have been
exhausted (Hegarty, 1977a).
Itnbibed seeds of lettuce held under induced thertno-
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
to 90% (Toole & Toole, 1953). In an extension ofthis
work (Villiers, 1974) seed viability in lettuce and ash
{Fraxinus americana L) was maintained over 12 months
in seeds kept fully imbibed, but was cotnpletely lost in
2 months (lettuce) and 3 months (ash) in seeds equili-
brated at 100% RM and then stored. Chromosotne

aberrations increased sharply in the vapour-equilibrated
seeds at increasing moisture contents but were absent in
the fully imbibed seeds, indicating an ability of tlie
dormant seeds to prevent storage deterioration. The
water content of the lettuce seeds equilibrated at 100%
RH was only 14% (dry weight or wet weight not indi-
cated) and Villiers speculated that a higher moisture con-
tent would be needed before macromolecular repair
and membrane synthesis could occur at a normal rate.
Dormancy can also be due to physical constraint of
the embryo by the seed coverings (Crocker, 1916;
Mayer & Poljakoff-Mayber, 1975). In this situation
embryo activation clearly occurs and the constraint
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-
George & Williams, 1968).
It has been implied that dormancy is an inactive
resting phase (Toole & Toole, 1953; Yemm, 1965).
Haber & Luippold (1960b) suggest that such an implica-
tion is quite inappropriate to describe the physiological
state of seeds prevented from germinating by with-
holding morphogenic stimuli. It is not generally related,
either, to a significant depression of overall metabolism
or to a block preventing cell division. Instead, they
suggest tliat it is a subtle block that specifically prevents
the initiation of cellular expansion. In many ways this
situation seems to be remarkably similar to seeds held at
the brink of germination by moisture stress, as in the
seed 'priming' treatment.
(7.4) The physiology of growing root eells
The subject of the transformation of meristematie into
elongating cells in roots has been reviewed by Obrou-
cheva (1975). Both transformation and elongation are
accompanied by acceleration of all metabolic processes.
The content per cell of numerous compounds is much
higher in elongating than in meristematie cells, both in
the case of compounds of primary metabolism as well as
the end products. Most compounds of primary metabol-
ism are osmotically active and tlie increases in their levels
in elongating cells favour decreased solute potential and
increased water uptake. These changes are accompanied
by parallel increases in enzyme activity and in respira-
tion, althougli the balance between the predominant
respiratory pathway through glycolysis and the Krebs
cycle on the one hand, and through the hexose mono-
phosphate pathway on t)ie other, remain constant. Thus
elongating cells represent sites of active metabolism of
a type that, as in meristematie cells, is dependent upon
the normal course of protein and nucleic acid syntheses
and is sensitive to inhibitors of those processes. How-
ever, cell elongation does not necessarily follow the
completion of cell divisions in the meristem, and
although the mechanism governing the process of trans-
formation itself and the initiation of elongation is not
known, it has been found to be insensitive to inhibitors
of protein and nucleic acid syntheses.
The dependence on different metabolic processes of
SEED HYDRATION AND DEHYDRATION 111
cell division and elongation on the one hand, and the
initiation of cell elongation on the other, may, if appli-
cable in seeds, be of fundamental importance in the
control of germination. If the water potential threshold
for the initiation of cell elongation is higher than that
for other processes of cell development (i.e. if the initia-
tion of elongation is the process most sensitive to
moisture stress) then at the appropriate water potential
seed development miglit be permitted but not active cell
elongation, and the initiation of cell elongation would
be the block in the germination process that allows seeds
to be brouglit to the 'brink' of germination but no
further.
(8) 'False' germination
Up to this point it has been loosely assumed that germi-
nation, identified as radicle growth from the seed,
represents a continuing phase associated with concurring
cell division and cell elongation. Rogan & Simon (1975)
have described an initial phase of slow elongation of
the radicle immediately after imbibition before a more
rapid phase commences and before mitosis is detected.
This phase of growth is therefore presumably associated
only with the elongation of existing cells in the embryo,
and Haber & Luippold (1960a) concluded that rootlet
protrusion in lettuce is a result of cell elongation, with
cell division playing no part. If conditions are such that
some cell elongation is permitted but no cell division,
than a 'false' germination may result. This has indeed
been described in, for instance, sorghum (Sorghutti
vulgare Pers.) (Nutile & Woodstock, 1967) and in some
grasses (Watt, 1974). In the latter case, radicle protru-
sion witliout further growth was a response to low water
potential, occurring between 0.5 and I.O MPa, and
appeared to be a resting phase before growth continued
in some seeds at the same water potential, or in all
seeds on transfer to free water. In one species in
particular, these 'hydropedetic' seeds were resistant to
dehydration injury and germinated readily on
rehydration in free water.
Hadas & Stibbe (1973), considering tliis phenomenon
in chickpea (Cicer arietitmtn L.) and maize seeds, suggest
that there exist critical hydration levels within the seed
which will permit various growth processes. The first
critical level permits cell elongation whereas the second
permits cell division in the radicle.
Whilst it is probably the initiation of cell elongation
that is sensitive to water stress in this instance, the idea
of differential sensitivity of different growth processes
to water stress is supported by the evidence that the
threshold water potential for shoot growth is higlier
than that for root growth (Dasberg, 1971; Gray &
Steckel, 1976). 'Sprouted' but non-emerged seeds were
found by Doneen &, MacGillivray (1943) in dry soils,
and these may represent 'false' germination, or radicle
growth without shoot growth as described by Dasberg
(1971). Restricted radicle growth without shoot growth
is also a phenomenon of low temperature (Kotowski,
1926) and this would coincide witli the finding of Haber
112 T. W. HEGARTY
& Luippold (1960a) for lettuce that at low temperature
mitosis is restricted more than cell elongation.
(9) Does the osmotic inhibition of germination resemble
a dormancy mechanism?
Evidence is accumulating that germination that is just
inhibited by osmotica such as polyetliylene glycol or by
NaCl can be restored to a greater or lesser extent by a
stimulus given to the seed directly or through the germina-
tion medium. The evidence is particularly clear in the case
of lettuce, in which osmotic inhibitionhas been overcome
by gibberellic acid added to the germination medium
(Kahn, 1960), by red light (Kahn, Goss & Smith, 1957;
Schiebe & Lang, 1965; Nabors & Lang, 1971) and by
growth regulators applied to the seed (Odegbaro &
Smith, 1969; Kaufmann & Ross, 1970; Braun & Khan,
1976; Khan & Knypl, 1977).
Whilst lettuce may not be typical in its response
there are, in addition, other reports of a similar nature in
other species. Osmotic inhibition in celery was overcome
by growth regulators applied to the seed (Klian &
Knypl, 1977) whilst partial release of salt effects using
growth regulators has been described for wheat (Chaud-
huri & Wiebe, 1968; Idris & Aslam, 1975), pea (;Uprety
& Sarin, 1973) and Spergularia media (L.) C. Presl.
(Ungar & Binet, 1975). In wheat, presoaks in some
growth regulators were more effective than water alone
in overcoming osmotic inhibition during subsequent
gennination (Darra et al., 1973). Finally, gennination in
the dark was higher than in the light in osmotically
stressed seeds of cucumber (Cucumis sativus L.), radish
and sunflower (Helianthus annuus L.) (MeDonough,
1967).
Although in many cases only a portion of the seed
population was released from the osmotic inhibition,
nevertheless there is a phenomenon common to all
these reports. Seeds were stimulated to germinate in
conditions in which, without the stimulus, they would
not have germinated even though the conditions of
water supply were the same. Thus there is a strong sug-
gestion that the failure to germinate under moisture
stress, in some species at least, represents a positive
response to the environmental conditions and is, in fact,
a form of secondary or induced dormancy. In many
respects this phenomenon resembles the responses of
some seeds to high temperature in which germination
inhibition can be overcome by treating the seeds directly
with growth regulators (Biddington & Thomas, 1976;
Heydecker & Joshua, 1976; Dunlap & Morgan, 1977).
(10) Ecological relevance
In a recent review Matthews (1976) proposed that seed
and seedling adaptations were ecologically significant
because they tended to minimize post-germination
seedling losses. He differentiated between adaptations
which influence the initiation of germination and those
which influence the chances of seedling survival, but
recognized both as being means to the same end, the
reduction of seedling mortality. He considered that
recent work on the initiation of germination has concen-
trated on dormancy mechanisms, both innate (primary)
and imposed (secondary). In general, dormancy is a
means by wliich seeds avoid germinating at a time when
seedlings would be placed in high-stress conditions.
The failure to germinate under moisture stress as des-
cribed in the previous section would constitute a simple
example of an imposed dormancy mechanism which in
many species (lettuce may be an exceptionKalm,
1960) is apparently readily reversible in an improved
water supply. This, together with the differential sen-
sitivities of radicle and shoot growth discussed in Section
8, would have obvious survival advantages in dry condi-
tions. Certainly, the ability of seeds to germinate at low
water potential can lead to establishment failures if dry
weather ensues, whereas seeds with more exacting re-
quirements for germination (often species native to semi-
arid areas) can establish more successfully (McGinnies,
I960; McWilliam & Phillips, 1971; Mott, 1974; Watt,
1974).
The first stage of imbibition is the result of the
physical properties of the seed (Mayer & Poljakoff-
Mayber, 1975), and Yamamoto (1964) has proposed
that the retention of absorbed water may be an adapta-
tion to xeric conditions. It might equally well be argued,
however, that rapid dehydration could prevent germina-
tion of seeds which, had they germinated, would have
exposed the seedlings to a higli moisture stress environ-
ment (Mott, 1974).
An alternative strategy promoting seedling survival in
drying conditions would be the ability for seedlings
after germination to grow at water potentials lower than
those permitting germination. Some evidence for this is
provided by the results of MeDonough (1975) who
showed that seeds removed from a water supply after
initial imbibition could continue to germinate, and that
equilibrium water potentials were eventually attained
which were lower than those that could have permitted
germination.
(11) Membrane effects
(11.1) Membrane integrity
As mentioned in Section 4.2, seeds soaked in water
imbibe, and can lose a wide range of solutes. The leach-
ing of solutes into the soak water has been used as a
seed vigour test in large-seeded legumes such as peas and
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
cotyledons showed the highest level of leaching but the
large range of levels found among seed lots resulted from
differences in leaching from live tissue (Matthews &
Rogerson, 1976). Whilst increased solute leakage may be
partly due to induced cotyledon damage in these large
seeds during imbibition (Pollock, Roos &Manalo, 1969;
Rowland & Gusta, 1977), membrane permeability ap-
pears to be the main property governing the extent to

which leakage will occur. This subject has been exten-
sively reviewed by Simon (1974) who considers that
leakage occurs until the phospholipid membranes of the
cells become intact, with this occurring most rapidly in
the outer layer of cells. Loss of seed vigour associated
with an inciease in seed leachate is put down to a de-
terioration of the lipid membrane. In addition, the
imbibitional cliilling injury found in seeds such as maize,
cotton, lima bean (Phaseolus Iwtatus L), French bean,
soybean and sorglium, which can be prevented if the
seed moisture content is raised before chilling (Cohn &
Obendorf, 1976), if the imbibition takes place for a
short period at a higiier temperatuie before chilling
(Pollock & Toole, 1966), or, in cotton, if the seeds are
given a hydration-dehydration treatment at higli tem-
perature before chilling (Thomas & Christiansen, 1971),
is also thought to be associated with membrane proper-
ties (Simon, 1974).
Bramlage, Leopold & Parrish (1977) have reported
that in soybean, humidifying the embryos to 25-30%
moisture (dry or wet weight basis not indicated) reduced
leakage and overcame cotyledon cracking during imbi-
bition, but did not eliminate continued leakage at low
temperature or imbibitional chilling injury, whereas in
embryos humidified to 35-50% moisture before imbibi-
tion, leakage at low temperature and imbibitional
chilling injury were eliminated.
From work on peas, Simon & Wiebe (1975) propose
that embryos with a moisture content of 43-54% (water
potentials of 8 to 5 MPa) will show httle or no
leakage during imbibition. The degree of hydration of
the membrane at these water potentials is not known,
but it is suggested that matric forces in the cells are
reduced to such an extent that water is available to stabi-
lise the membrane bilayer. This level of water availability
is sufficient to permit the formation of ice crystals on
freezing.
(11.2) Other membrane properties
Leakage of salts from seeds has also been reported for
radish (Cocucci & Cocucci, 1977) but in this case the
leakage was followed by active salt uptake, at least in
the case of K'^. Addition of abscisic acid (which inliibits
radish germination) to the medium enhanced the salt
loss whereas fusicoccin (which stimulates the rate of
germination) promoted rapid, active K'^ uptake. The
authors suggest that activation at the cell membrane
level of some metabolically-dependent mechanism of
K* uptake probably linked to H"^ release is associated
with germination in radish seeds. Fusicoccin probably
enhances K* uptake capacity whereas abscisic acid in-
liibits it, and germinatioii could be linked with H*
weakening of the cell wall and/or K"^ uptake decreasing
the solute potential of the cell, both mechanisms en-
couraging water uptake.
A general role of membranes in the control of germi-
nation was suggested by Mayer & Sliain (1974) on evi-
dence from several sources. The involvement of the
membrane in the specific control of solute transport.
SEED HYDRATION AND DEHYDRATION 113
though not related to germination in particular, and the
effect on this of growtli regulators acting via the mem-
brane, have been reviewed by several contributors in the
volume by Marre & Ciferri (1977). In addition, Cocucci
& Cocucci (1977) point out that fusicoccin appears to
be especially effective in overcoming germination
inliibition in lettuce resulting from moisture stress or
liigh temperature (Braun & Khan, 1976), thus again
suggesting control over transport relationships.
The potential importance and involvement of the
cell membrane has been further indicated by the possi-
bility that pressure itself may control transport processes
in the plasmalemma and tonoplast. Coster, Steudle &
Zimmermann (1976) suggest that the cell membrane is
compressible by both mechanical and electrical forces,
and that significant changes in the tliickness of the
membrane can occur as a result of direct compression
due to liigli turgor pressure, indirect effects due to
stretching of the cell wall, and the stresses induced by
the electric field in the membrane. They suggest that
changes in membrane thickness could provide the
pressure-transducing mechanism required for osmo-
regulation and growth. Hence the rate of pumping of
ions (and thus the solute potential of the cell sap) could
be directly controlled by the turgor pressure.
(12) Germination control
There is a major problem in bringing together informa-
tion from widely different species involving different
seed structures, with the accompanying different re-
lationships between the embryo and its enclosed or
surrounding reserve material, different seed constituents
and different dormancy types and levels, in which ger-
mination control mechanisms may be entirely different.
A large amount of work has been carried out on the
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
germination control resulting from a balance of growth
regulators within the seed (Khan, 1975), with germina-
tion probably operating through a matrix of interacting
metabolic pathways and with growth regulators influ-
encing the enzyme activities within this matrix (Hey-
decker & Coolbear, 1977). Added to this is the problem
that release of germination inhibition by stimuli (as
opposed to the leaching of an inhibitor) may act by
different means. In the case of lettuce, for instance, it
has been suggested (Schiebe & Lang, 1965) that the
dormancy caused by embryo constraint within the endo-
sperm can be broken by irradiation in red light, inducing
greater embryo thrust, or by gibberellin which possibly
acts by weakening the endosperm.
There are, however, some significant features that
can be extracted from the data wliich may point to an
understanding of germination control and the effects of
seed hydration treatments.
(1) In the work just mentioned (Schiebe & Lang,
1965) it was found that red hght caused increased em-
bryo growth of lettuce seeds in mannitol solution. This
114 T. W. HEGARTY
work has now been taken further to show that phyto-
chrome stimulation of lettuce axes held in polyethylene
glycol solution can result in a decrease of solute poten-
tial within the embryo of 0.1-0.2 MPa, and a change in
pressure potential of a similar order such that in total
the embryo overcomes the constraint of the endosperm
(Carpita & Nabors, 1977). According to Black (1969)
seeds must be undergoing active respiration in order for
phytochrome to act. The point is that given certain
external conditions, osmoregulation within the seed was
clearly affected by the external stimulus.
(2) Pressure can affect cell membrane properties and,
through this, osmoregulation (Coster et al., 1976).
Changes in water potential could well affect cell mem-
brane properties either by acting directly on the mem-
brane, or indirectly by affecting the pressure relations of
the cell.
(3) Growth regulators could simOarly act directly on
ion transport processes in the cell membrane (as sug-
gested by Marre (1977) for fusicoccin), or indirectly
through cell metabolism but ultimately affecting trans-
port processes in the membrane (e.g. weakening of the
cell wall). Braun & Khan (1976) point out that effects
of water stress have been associated with alterations in
the levels of endogenous growth regulators and, in seeds,
exogenously applied growth regulators can sometimes
break dormancy, including that induced by water stress.
Clearly, this action could be through an effect on the
cell membrane and solute transport.
(4) Some cell division may apparently occur in the
embryo at water potentials lower than those that permit
the co-ordinated division and extension associated with
continuing radicle growth. If the situation described in
roots by Obroucheva (1975) can be applied to the ger-
minating seed, then it may well be the process of the
initiation of elongation, which apparently has quite
distinct metabolic properties, which is involved in the
control of radicle emergence and its continuing growth.
This process is presumably concemed with osmoregula-
tion within the elongating cell. The work of Fujisawa
(1966) who showed that phase C growth of radish axes
could be inhibited without interfering with phase B
respiration, miglit also suggest that it was the initiation
of cell elongation that was being inhibited.
(5) The proposal that the transformation of cells to
the elongating stage has a higher water potential threshold
than embryo development or growth, implies as a conse-
quence that germination once initiated could proceed at
lower water potentials. This is supported by the findings
of MeDonougli (1975, 1976) that germinated seedlings
do develop low water potentials.
(6) In some cases, imbibition under unfavourable con-
ditions affects subsequent germination, presumably
through an effect on the hydration of membranes
(Simon, 1974) and initial, even partial, hydration under
more favourable conditions can overcome this problem.
Seed ageing results in membrane deterioration and a
reduction in the range of conditions in which seeds will
germinate, and it is possible that in low-vigour seeds of
this category, membrane hydration or integrity effects
are at least partially responsible for determining the
success or failure of germination.
(7) The fact that in a population of seeds different
numbers germinate over a range of water potentials,
and that pre-soaking treatments can alter these responses,
imphes that seeds in a population have a range of
threshold levels for initiation of germination and that
the threshold levels can be altered. This could be inter-
preted as a membrane effect (e.g. allowing hydration or
even repair under favourable conditions) but in any case
it is evidence that events occurring later in germination
are dependent upon events occurring earlier in the germi-
nation process.
(8) Some seeds can imbibe and be held at low tem-
peratures for very long periods yet still germinate nor-
mally on transfer to favourable conditions (Harrington,
1962). In a species such as carrot, hydration treatments
at temperatures favourable for germination can help to
overcome low-temperature sensitivity (Hegarty, 1970)
and presumably in these cases membrane effects as
described in paragraph 6 above are not involved. Austin
et al. (1969) have suggested that in such cases hydration
treatment may allow the embryos of some seeds to
'after-ripen' and in this case it may be chemical or mor-
phological, or possibly membrane, modifications that
permit some seeds to germinate at low temperature that
would not have germinated without the hydration
treatment.
(9) Germination itself must be associated with the
osmotic regulation of cells in the radicle. Such osmotic
regulation can apparently be influenced in a number of
ways in seeds. It seems likely that many, if not most of
the processes that occur in seeds supplied with water
sufficient to germinate also occur in seeds held under
moisture stress such as that of the seed 'priming' treat-
ment (though perhaps more slowly), the main differ-
ences being associated with the process that induces cell
elongation. It may well be significant that this process
could be under a metabolic control that apparently does
not infiuence other processes within the seed associated
with preparation for germination (i.e. phases A and B of
germination).
(10) There is a strong resemblance between the
'subtle' block to germination described by Haber &
Luippold (1960b) for some types of dormancy and the
block to germination of seeds held under moisture stress.
(13) Conclusions
Initial seed imbibition is apparently a physical process.
Seeds in contact with a free water supply take up more
water than they need to germinate, though the absolute
quantity taken up will depend on the seed structure and
constituents, on the temperature and possibly on other
properties of the external environment. At lower levels
of water potential seeds imbibe less and germinate more
slowly until eventually germination is prevented. This
varies between individual seeds within a seed lot, be-
tween seed lots of the same species, and between species.
Even when seeds are prevented from germination by

the water potential of the external etivironment, it is
abundantly clear that there is a range of water potential
within which many, if not all, of the processes leading to
normal germinatioji up to a particular point can occur,
and that substrates for future growth may accumulate,
or systetn activity may be enhanced. Several aspects of
the environment appear to affect the precise extent to
which all the seeds within a seed lot may be brought to
the brink of germination. Seeds unable to germinate for
other reasons (various types of dormancy) have also
been shown to be metabohcally active, and sotne forms
of dortTiancy show similarities to the inhibition of germi-
nation by moisture stress.
Dehydration before germination tnay suspend activity
and development at the level just prior to dehydration,
but may also be associated with gross or more stibtle
forms of embryo damage (includitig damage to tlie cell
membrane), or with the induction of fotms of secondary
dormancy.
One can speculate that the process in the embryo
most sensitive to moisture stress is the initiation of cell
elongation. This is likely to be associated with osmo-
regulation within the cells. It appears that osmoregula-
tion is sensitive to various factors within the embryo
which tnay act independently or through a common
pathway, and mucli evidence suggests that the cell
metnbrane is actively involved. Osmoregulation is sensi-
tive to external stimuli and this suggests that the failure
to germinate under moisture stress can, in some cases at
least, be a positive response to the environment, and can
be classed as a form of secondary dormancy. In roots,
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
applicable in seeds, this could give the advantage of
allowing normal embryo development to occur in
drought-stressed seeds whilst preventing germination.
The phenomenon of physiological development of
the embryo without germination occurs in natural habi-
tats and, under some circumstances at least, may lead to
enhanced seedling sumval. Conversely, sowing pre-
germinated seeds may place the seeds seriously at risk in
the environment by removing gertnination control from
the seeds' environment-sensitive mechanisms.
In some species it is thought that the conditions of
initial seed hydration affect membrane integrity and
hence the subsequent ability of the seed to germiriate
normally. Such conditions include, for some species,
low-temperature imbibition or seed soaking. Seeds that
are insufficiently hydrated deteriorate, and two phe-
nomena associated with loss of vigour through seed
ageing are membrane deteriotation and a reduction in
the range of conditions over which seeds can germinate.
It is thus possible that membrane hydration effects may
be at least partly responsible for restriction of the range
of gertnination conditions associated with seed ageing.
At progressively higher equilibrium moisture con-
tents, the rates of seed metabolism and deterioration
also increase compared to dry seeds. However, if the
equilibrium moisture content is sufficiently higli, seed
SEED HYDRATION AND DEHYDRATION 115
activation or repair processes counteract or replace the
mechanisms of seed deterioration. A limited atnount of
work suggests that membrane integrity is not achieved
until water potentials of 8 to S MPa are attained, not
very much less than the levels (down to - 2 to - 4 MPa)
usitig osmotica or vapour treattnents at which known
benefits to seeds have been reported. It is possible that
seed activation or repair mechanisms cannot operate
either effectively or at all until tnembrane integrity has
been attained, and that deterioration mechanisms domi-
nate or operate alone at hydration levels below these
water potentials (8 to 5 MPa). Tliese levels are in
equilibrium with very higli RHs (95% +) and the prob-
letns associated with maintaining accurately such RHs
could well explain at least some of the conflicting
results obtained with seeds held in so-called saturated
atmospheres.
The likelihood of finding common trtechanisms of
gertnination cotitrol and seed activation or deterioration
under moisture stress seems in many ways highly un-
likely because it is diversity that encourages survival.
Nevertheless, whilst speculative, some of the suggestions
above do offer links between the findings of many ex-
perimenters working with quite different objectives in
mind, and several lines of work suggest that the cell
tnetnbratie tnay be futidatnentally involved. Tliis hy-
bridisation of ideas may stimulate a number of critical
experiments to be performed which, at very least, should
yield information on the physiology of incompletely
hydrated seeds, a field in which very little information is
currently available.
Acknowledgments
I am indebted to Dr John Raven of the Department of
Biological Science, University of Dundee, for helpful
discussion during the preparation of tliis paper, including
the indication of a nutnber of relevant references, and to
Dr Peter Waister of the Scottish Horticultural Research
Institute for useful comments on the content and
construction of the tnanuscript.
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