SEED DORMANCY

Seed dormancy is the physiological state in which a viable seed does not germinate , even in
the presence of favourable environmental conditions. Dormancy is a device that optimize the
distribution of germination in time or space, and it has great ecological importance. This is
possible because seeds of many species show variability in depth of dormancy, which exhibits
sporadic release from dormancy resulting in irregular germination. The temporal dispersal thus,
enhances the spread and survival of the species. Seasonal environmental changes (light and
temperature) ultimately regulate seed dormancy and germination, controlling the seed dispersal.

Four basic patterns of germination have been recognized in relation to its temporal
distribution.

1. Quasi-simultaneous- when germination of all the seeds occurs over a relatively brief period.

2. Continuous- in which seeds germinate over an extended period of time. 

3. Intermittent- irregular germination over long periods of time (multimodal distribution) 

4. Periodic- multimodal germination showing more regular Periodicity.

These patterns result from the dormancy characteristics and from their interplay with
various environmental factors like light and temperature. Dormancy is also important for
preventing vivipary and precocious germination i.e., germination while the seed is on the mother
plant, which is important in certain crop species like cereals.

Dormancy is most commonly found in species which occur in areas of seasonally harsh climates,
and enables the organism to survive as a seed when it might perish as a seedling eg., during a
cold winter or a long dry season. Even in moist tropical forests, dormancy occurs in pioneer,
light demanding, gap-colonizing genera such as Macaranga, Trena, etc. Seedlings of these genera
are incapable of surviving under the low light conditions found under undisturbed forest canopy.
Dormancy is useful in enabling the species to survive as seeds until a gap is formed by natural or
man-made causes. Dormancy is broken when the seeds are exposed to the increased light and
temperature in canopy gaps. In drier tropics, seed coat dormancy is common, most probably a
built-in mechanism to withstand unfavourable environmental conditions. In nature, the dormancy
or after-ripening phenomenon lasts till the conditions for germination become favourable. Seeds
of tropical hard woods in central India for example, mature during summer months; in case these
seeds did not possess the after ripening phenomenon (post harvest dormancy) the young
germinants would not have survived eg., Tectona grandis.

Classifications

  Seed dormancy can be classified based on

  i) Manner of origination (Harper, 1977)

1. Innate dormancy 

2. Induced dormancy

3. Enforced dormancy

  ii) Time of origination

1. Primary dormancy

2. Secondary dormancy 

iii) Location of dormancy

1. Exogenous dormancy (Seed coat dormancy) 

2. Endogenous dormancy (Embryo dormancy)

3. Combined dormancy

  iv) Cause of dormancy

1. Physical dormancy

 2. Chemical dormancy 

3. Mechanical dormancy 

4. Morphological dormancy 

5. Physiological dormancy 

6. Morpho-physiological dormancy 

7. Combinational dormancy 6 and 7 are called Double dormancy

A summary of dormancy types according to the classification is discussed as follows.

Innate dormancy

This type of dormancy is present when the seeds are ready for dispersal. lt prevents
germination during development and maturation of the seed on the mother plant and also
for sometime after shedding and harvesting. Seeds with impermeable coats and those requiring
prechilling treatments represent innate dormancy. This is also referred to as primary dormancy.
Induced dormancy

A seed may become dormant if, during germination, it is exposed to unfavourable conditions
when it is predisposed to germinate viz., excessive moisture or desiccation, extremes of climates
etc. This is also referred to as secondary dormancy.

Enforced dormancy

When a light requiring seed is buried deep in soil and fails to germinate due to non-availability
of light, this is called as enforced dormancy.

Exogenous dormancy

Dormancy caused due to factors relating to seed coat or pericarp. Analogously, mechanical
resistance, physical impermeability, inhibitors or light sensitivity are associated with the seed
coat.

Endogenous dormancy

Dormancy caused due to factors relating to embryo eg., immature development or chemical
inhibitors located in the embryo.

Combined dormancy

Where two or more types occur in the same seeds it is called combined dormancy. Dormancy
may be caused due to factors relating to seed coat as well as embryo.

A comprehensive classification

A simplified classification and terminology is presented hereunder which basically follows one
used by Nikolaeva (1977)and Baskin and Baskin (1998) with few modifications.

A. Exogenous dormancy

i) Physical : Impermeability of seed coat or pericarp to water

ii) Chemical : Inhibitors in pericarp or seed coat

iii) Mechanical : Mechanical resistance of pericarp on seed coat

to embryo growth.

B. Endogenous dormancy

i) Morphological : Under developed embryo

ii) Physiological : Physiological inhibiting mechanism

a) Shallow : Weak inhibition

b) Intermediate : Intermediate inhibition

c) Deep : Strong inhibition.

Thermodormacy: Temperature sensitive embryo

Photodormancy : Light sensitive embryo

C. Combined morpho-physiological dormancy

Combination of under-developed embryo with strong physiological inhibiting mechanism of

epicotyl growth

D. Combined exogenous / endogenous dormancy (A+B)

A. Exogenous dormancy

1. Physical dormancy

This is caused due to seed coats impermeable to water. The impermeability is due to a layer of
thick walled, cutinised paliside like macroscleroid cells. If the coat of such seeds is ruptured or
softened, so as to permit the ingress of water, the state of dormancy may be overcome eg.,
Acacia nilotica and Albizzia lebbeck.

2. Chemical dormancy

Seed coats may contain germination inhibitors. These seeds lack physiological dormancy and
thus the inhibitor chemical can be leached out of the seed or some how deactivated. Chemical
dormancy is broken by removal of pericarp or leaching of the fruits. Light sensitive inhibitors are
present in intact seeds of Betula pubescens and Betula pendula.

3. Mechanical dormancy

It is due to the presence of a hard woody fruit wall usually endocarp some time mesocarp. It is
proposed that the endocarp acts as a mechanical obstacle to a germinating embryo.

In general, once the seed absorbs water and if the embryo is not dormant, the force of
expansion of the germinating embryo ruptures the seed coat and breaks apart the coverings. But,
the coats of many seeds and indehiscent fruit eg., nuts, are hard and have very tough tissues,
although they allow entry of water, yet offer resistance to the growth of germinating embryo.
This is because the internal pressure developed is insufficient to burst the seed, a process that
must be effected before swelling can be completed and germination commenced. eg., Eucalyptus
delegatensis and E. pauciflora.

Long periods of cold stratification are required for loss of de mancy. Once the dormancy of
embryo is broken however, it ha growth potential to push through the endocarp. Thus it appears.
mechanical dormancy in these species should be reviewed as an aspect or manifestation of
physiological dormancy.

B. Endogenous (embryo) dormancy

1. Morphological dormancy

In some species, at the time of dispersal the embryo is not fully grown to get differentiated into
radical and cotyledons, there fore ger mination does not take place until both differentiation and
growth occur. Fraxinus spp, requires about 4 months to complete development: In case of
Ginkgo biloba even fertilization of the ovule is said to take place after the seed falls.
Requirements for growth are moist substrate, suitable temperature and specific dark : light
condition. Gibbrellic acid also increased germination of certain species at super optimum
temperature.

2. Epicotyl dormancy

In seeds of some plants, only a part of the embryonic axis may be dormant. In Viburnum spp, and
Asarum canedense, the radicle growth proceeds normally while the embryo and embryonic axis
remains dormant at room temperature. The hypocotyl or root of such seeds elongate and develop
until the stored food is consumed, after which it withers and dies. This is called epicotyl
dormancy, which may be overcome by chilling or other dormancy breaking treatments. Exposure
of such germinated seeds to low temperatures (33 - 50° F), may induce the tops to resume
growth.

3. Physiological dormancy

A. Non Deep Physiological dormancy

Freshly matured seeds with non deep physiological dormancy either cannot germinate at any
temprature (or) they germinate only over a very narrow range of temperatures (Thermo
dormancy) eg., Fagus, Pinus and Eucalyptus. Such dormancy in some species is broken by
relatively short periods of cold stratification ranging from 5 days in Triticum sp to 60-90 days in
Impatiens biflora. Seeds stored dry at room temperatures come out of dormancy after ripen.
However, the time required for seeds to after ripen is usually much longer than that required for
dormancy loss during cold stratification. Dormancy loss may occur at high temperatures
regardless of whether seeds are imbibed. Non deep physiological dormancy can be broken by
chemcials, including potassium nitrate, thiourea, kinetines, ethylene and gibbrellin.

A light requirement for germination is another manifestation of non deep physicological

dormacy (Photo dormancy) eg., Spathodea and some Eucalyptus. Seeds of some species lose
their requirement for germination as they come out of dormancy in response to cold stratification
or high summer temperatures, but some species have a light requirement for germination after
they have recieved dormancy breaking treatments.

Physiological dormancy is caused by embryo and by interaction between the embryo and
its covering structures,

i) Seed coat may restrict movement of oxygen to the embryo or inhibitors (eg., phenolics in seed
coat may fix oxygen by oxidation, thus making it unavailable to embryo).

ii) Seed coat may prevent leaching of inhibitors, retard the entrance of the oxygen which can
inactivate or prevent production of inhibitors or themselves contain growth inhibitors. After
breaking of dormancy, embryo becomes insensitive to inhibitors, even though it is present.

iii) The embryo covers i.e. the seed coat may mechanically restrict embryo growth. If seeds are
placed under appropriate dormancy breaking and / or germination conditions, the growth
potential of the embryo increases and germination occurs. Depending on the species, cold
stratification, gibbrellin incubation temperatures, light or dark may increase the growth potential
of the embryo.

iv) Endosperm rather than seed coats are the main force restricting embryo growth and thus
germination in some species. Resistance has been shown to vary depending on the temperature
and dark : light conditions under which seeds were imbibed and was also influenced by
gibberellins.

v) In many species interaction between embryo and covering structures can be the best
explanation for cause of dormancy. Embryo may produce signal to stimulate production of
hydrolases in endosperm. Embryo may remove or absorb products of enzyme hydrolysis that
inhibit enzyme activity. Embryo may also regulate inhibitors in the endosperm.

B. Intermediate Physiological Dormancy

Embryos isolated from the seeds with intermediate PD will and the resulting seedlings are
normal. The dormancy of intact or dispersal units is broken by cold stratication but upto 6
months this treatment may be required.

C. Deep Physiological Dormancy

Embryos isolated from seeds with deep PD either do not grow or they produce abnormal
seedlings. The only treatment that overcomes the dormancy of intact seeds (or dispersal units) is
a relatively long period of cold stratification.

D. After-ripening

In this type, no morphological changes can be seen from the time the seed falls from the tree,
until it is fully developed and is ready for normal germination. However, the physiological
process that intervenes is called "after-ripening". This proceeds most actively at a low
temperature (6-10°C). 

Chemical changes accompanying after- ripening are listed as follows: 

1. Softening of the seed coat 

2. Increase in amount of water absorbed

Increase in total acidity. Usually, the hypocotyl is at first alkaline but becomes acidic when the
after-ripening is complete, the H-ion

concentration and enzyme activity also increase.

 3. Treating with dilute acid (HCL, acetic acid, butyric acid) may shorten

the period required for after - ripening. The increase in acidity is

often correlated with greater water absorbing power of colloids 

4. Increase of reducing sugars, amino nitrogen and amino acids 

5. Increased respiration 

6. Appearance of HCN in cotyledons 

7. Increase in catalase, oxidase and peroxidase activity 

8. Short chain fatty acids such as volatile fatty acids may induce seed dormancy and their loss
during after ripening may trigger dormancy termination.

Freshly collected seeds of Tectona grandis would not germinate, no matter how elaborate
weathering the seed has been subjected to. Other examples are Cassia fistula (12 months),
Samania saman (13 months) and Lagerstroemia flosreginae (7 months). Dalbergia latifolia,
Pterocarpus marsupium, Adina cordifolia, Albizzia lebbek, Cassia siamea, Terminalia chebula,
Tectona grandis, Zizyphus jujuba, etc., also require some period of after ripening.

C. Double dormancy (or) combined dormancy

In some species, seed dormancy is caused by the combined effect of factors associated with the
embryo and seed coat. Seed of Fraxinus excelsior exhibited deep dormancy, which was caused
by combined effect of restriction of oxygen supply to embryo and a requirement for chilling.
Therefore, more than one treatment may be necessary in order to break the double dormancy of
such seeds.

Treatments to overcome dormancy

Pretreatment for dormant seeds is a must to enchance rapid and uniform germination of seed
sown in the nursery, field or during seed testing.

i) Mechanical scarification

Large quantities of hard coated seeds can be mechanically scarified using concrete mixer with
sharp gravel or sand; special drum lined with abrasive material such as sand paper, cement or
crushed glass or fitted with abrasive disks may be made. Care should be taken to avoid over
treatment eg., Acacia catechu, A. nilotica, Albizzia falcataria, A. lebbek, Cassia fistula,
C.javanica, C. nodosa, Delonix regia, Santalum album, Terminalia arjuna, T. tomentosa.

ii) Water soaking

This treatment is intended both to soften hard seeds and to leach out chemical inhibitors. Soaking
in cold water for a period ranging from a few hours to several days, is a safe and simple
treatment, which can be tried on all, but the smallest seed. Leguminous seeds often respond well
to immersion in boiling water; the seed to be treated being placed in a quantity of boiling water
more or less equal to its own volume and left till the water cools. eg., seeds of oaks and chestnuts
are reported to respond well to a 1/2 minute immersion in boiling water. Soaking in water for
periods ranging from 2 to 48 hours according to species viz., Acacia mearnsii, A. nilotica,
Adenanthera microsperma, Albizzia amara, Grevillea robusta, etc.Prescriptions for hot water
treatment must be applied meticulously to remove seed coat dormancy without killing the seeds
through successive heating.

iii) Acid scarification

Soaking in concentrated sulphuric acid is the most common method of treatment to soften the
seed coat. For this, the material needed include commercial grade Sulphuric acid (95%, 36N),
acid resistant containers, wire containers, screens and an abundant supply of water for rinsing the
seeds after treatment. To scarify the seeds, pour the undiluted acid until all the seeds are wetted.
Stir them for few minutes to one hour depending on the species and immediately wash them
thoroughly in cool running water for 5-10 minutes to remove all traces of acid. Shade dry the
seeds unless wet sowing is preferred. In seed coat of Prosopis juliflora, an oily deposit in the
coat is responsible for delayed germination, seeds germinated when this is dissolved in ether.

iv) Dry heat treatment

Spread the light inflammable litter over the fruits and then ignite it. However, adjusting the heat
of the fire to achieve the maximum effect on the pericarp without damaging the seed embryo
requires experience. The exposure of seeds of Acacia mangium to dry heat at 100°C was nearly
as effective as immersion in water at 100°C in overcoming dormancy. In Tectona grandis, the
drupes exposed to 45°C - 50°C showed 50 per cent germination.

v) Warm stratification
This has been recommended both to overcome mechanical and morphological dormancy. Soak
the seeds in several times their volume of cold water at approximately 3-5°C for 48 h. Drain off
the water and mix the seeds with two to four times their volume of a moistened, water retained
medium such as sand, sand peat mixture or vermiculite. Store at a warm temperature. A constant
temperature of 20-25°C or alternating temperature of 20°C and 30°C is suitable for many
species. Open the containers weekly, mix seeds and if surfaces show signs of drying out, remoist
with water spray.

vi) Cold stratification

This is recommended to remove physiological dormancy. Seeds should be soaked in

several times their volume of water before prechilling, a 48 h soak at 3-5°C. After soaking, the
water is drained off and moist seeds are stored in 3-5°C for the period appropriate to species.
Storage may be without any medium i.e. seeds may he mixed with 2-4 times its volume of a
medium such as moist sand, peat or a mixture of the two. Containers shall be opened and mixed
if surfaces dry off eg., Abeis spp, Eucalyptus delegatensis etc.
The combination of high moisture and low temperature appears to trigger off changes in
cell energy relationships leading to biochemical activity which transforms complex food
substances into s utilized by the embryo when it renews growth at germination. Low temperature
not only favours these biochemical changes, but also reduces microorganisms' activity and the
risk of overheating and premature germination of the after-ripened seeds.

vii) Biological methods

In nature, animals and microorganisms are an important factor in the break down of seed coat
impermeability. It is difficult to make use of these organisms as a controlled pretreatment of
seed, but in few cases successful results have been obtained. There is no doubt that certain seeds
germinate more freely after they have passed through the bodies of animals. The combination of
moisture, warmth and chemical action of the digestive juices doubtlessly softens and renders the
hard seed coat permeable. This method has been experimented with Santalum album, Acacia
nilotica and Macaranga denticulata, but effective control is very difficult.

Hormonal regulation of Seed Dormancy

1. Abscisic acid inhibits germination of non dormant seeds. It is the main chemical
factor for induction of dormancy frequently accumulating in maturing seeds.
2. Gibberellin stimulates seed germination of many species. Dormancies with
chilling and light requirements are often overcome by treating with GA
3. Cytokinin is more effective than GA in overcoming the effect of inhibitors like
ABA of various GA sensitive processes.
4. Auxin (IAA) may be associated with the later stages of embryo expansion
during seed germination, but may not play any significant role in development or
termination of seed dormancy in forest seeds.
5. Ethylene breaks the dormancy of seeds of many species and has been
implicated in the regulation of seed dormancy.