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During the developmental cycle of the plant, at some phase or the other certain structures like buds,
tubers, seeds, etc., go through a period of temporary suspension of growth activity. Such a state is
called dormancy. It may be imposed by certain environmental factors or internal factors or genetic
factors included. Generally plants or plant structures, in order to overcome or survive against hostile
environmental conditions undergo a period of dormancy with suitable modifications. In lower plants
production of endospores, zygospores, auxospores, akinetes, etc, are some of the methods involved
in tiding over unfavorable conditions. Even vascular plants with their complex structural
organization and reproductive methods produce dormant structures to overcome unfavorable
conditions. Some of the dormant structures that develop in plants greatly facilitate in the dispersal
mechanism. Among the many structures which exhibit dormancy, seeds and buds are important.
Bud Dormancy
Perennial plants like shrubs, trees have to go through different seasons in a year. The onset of
winter is always an unfavorable season for the growth and even survival of plants become difficult
because of extreme variations in the temperature, especially cold conditions. In order to survive
against such hostile conditions the growth regions like apical buds, axillary buds and underground
structures like rhizomes, tubers, etc, undergo a period of dormancy suspended growth. It is during
this season that leaves wither. In fact, it is not the change in the temperature that acts as the signal
but it is the change in the photoperiodic conditions which triggers the falling of leaves.
Aesculus hippocastanum-floral dormancy
Dormant cabbage
Quercus palustris
Protea dormant floral structures
It is again during the same season, meristematic cells found in the buds undergo temporary
suspension of their activities. In addition to it, the meristematic regions will be armed and protected
with a number of leaves called bud scales which are thick, waxy and covered with a dense coat of
hairs. Such structures provide thermal insulation to the meristematic zones and prevent them from
cold injury or frost bite.
Site of Perception:
Though leaves perceive changes in photoperiodic effects of the day, it is the buds that act as the
sites of perception for inducing dormancy. The induction of dormancy in buds starts only after the
falling of leaves. Nonetheless, in many plants even old leaves act as the sites of perception in
inducing bud dormancy. Such buds can be induced to break the dormancy by subjecting the same to
long photoperiodic treatment or interrupting the long dark periods by red light.
The onset of short day or long dark photoperiods in winter stimulates the synthesis of various
growth inhibiting compounds of which Abscissin dominates. Quantitative estimation by solvent
extraction methods reveal that dormant buds contain greater amounts of ABA than actively growing
buds. Abscissin is a well known growth inhibiting hormone. By inhibiting the synthesis of proteins,
RNA and other metabolic processes (See ABA chapter), ABA imposes dormancy on meristematic
tissues of the plant body.
The involvement of photoperiodic effect on bud dormancy indicates that the phytochromes have a
role in imposing dormancy. Phytochromes are known to be present in plastid membranes and other
surface membranes. It is also known that ABA is synthesized and often stored in
plastids. Phytochromes by perceiving the changes in photoperiodic conditions probably induce the
synthesis of ABA and also facilitate the release of ABA. Then ABA is translocated to the buds where
it brings about temporary suspension of metabolic activities and totally inhibits mitotic activity. The
presence of Abscissin II, which is also called Dormin has been detected from various plant
structures involved in dormancy ex., dormant potato tubers, birch leaves, winter buds in
clerodendron, pinus, etc.
The dormant buds can be induced to sprout again by treating with cytokinins and gibberellins. But in
natural course, the onset of spring and long photoperiods, the dormant buds become active and
develop into branches.
Cytokinins are known to be synthesized in root tips but under cold conditions because of the snow
fall, the root meristems are very inactive and they dont synthesize sufficient quantities of cytokinin
required for the buds to be active. That is probably one of the reasons why buds remain
dormant. As soon as cytokinins are provided to dormant buds, mitotic activity is initiated and buds
start sprouting. Besides, cytokinins also counteract ABAs inhibitory effect of the metabolic activity
level and promote growth activity.
Another class of phytohormones, which over comes the bud dormancy is Gibberellins. Now it is
certain that Gibberellin synthesis takes place in plastids. Moreover, the synthesis of GA and ABA
starts from the same precursor called mevolonate. Under short day conditions, the pathway from
mevolonate is directed towards ABA synthesis and GA synthesis is inhibited, but during long day
photoperiods it is directed towards GA synthesis and ABA synthesis is blocked. That is the reason
why gibberellins under long day conditions or not light treatment, break bud dormancy and nullify
the effect of ABA present in such dormant buds.
The photoperiodic effect either in breaking the dormancy or induction of dormancy is explained on
the basis of phytochrome involvement. When long day conditions prevail, more amount of PfR form
of phytochrome accumulates in the cells which initiates not only the synthesis of more GA and but
also it facilitates the release of it from the plastids into the cytoplasm. Once GA is released, it brings
about the activation of dormant cells, and thus GA breaks bud dormancy. It is really very fascinating
to understand the three way interaction between phytochrome, photoperiod and GA/ABA synthesis
in imposing bud dormancy or breaking it.
Seed Dormancy
Angiosperms produce seeds within the ovary and ovary itself develops into a fruit. In Gymnosperms
only seeds develop, because the ovule is naked and it is not enclosed by any ovary wall as in the case
of angiosperms. During the development of a seed or the fruit, some remarkable changes occur in
the ovule as well as in the ovary wall. The development of ovary into fruit and seed go hand in
hand.
Cotton seed
Sometimes, one develops faster than the other, but ultimately the slower one catches up with the
other at the time where fruit is ready to be shed.
Gene expression during dormancy and breaking dormancy
Left) Seeds of Arabidopsis thaliana Cvi that have lost dormancy (G) are characterised by low
expression of two dormancy genes (ATS2, ATS4) and high expression of two germination genes
(AtRPL36B, AtRPL27B). The two dormancy genes (ATS2, ATS4) display high expression in dormant
seeds (D) that do not complete germination. (Contact: Dr Peter Toorop
The duration of dormancy in seeds varies from species to species and it is species specific. It has
been noted that the duration of dormancy varies from few months to many years. The controlling
factors that impose dormancy are many ex., the strength or the susceptibility of seed coat, ability of
the seed to absorb and utilize water or oxygen, presence of inhibitors and environmental factors like
temperature, water, day length, etc.
Viability of Seeds:
The period for which the embryo remains healthy within the seed coat and capable of germination
under permissible conditions is referred to as seed viability. In spite of favourable conditions, seeds
fail to germinate beyond the viable period. In fact, the period of viability refers to the longevity of
the embryo. The half life of seeds vary from species to species, where some have few days or
months and some remain viable for a period as long as 100 to 1000 years. The conditions at which
the seeds are preserved are also important. Horace Wester (1971) reported that the lotus seeds
obtained at an excavation point were as old as 900-1200 years, still they were viable. Majority of
crop plants that are invaluable for human beings are viable for 1 to 2 years.
Seed Coat:
Many seeds are incapable of germinating immediately because of the hard seed coat which is
thought to break open by the developing embryo. It often does not imbibe water; even O2 does not
difuse in which are the most important factors favourable for germination. Thus the seed coat,
though it acts as a protective structure often imposes dormancy in many seeds in the following
ways. It is a necessary evil.
(1) Prevents water uptake: Some plants produce seeds with hard seed coat which is also
waxy. Thus the seeds are rendered impermeable to water. For example, in some Fabaceae
members like is Pinus arborous, water first enters through hilum which is made of hygroscopic
tissue. On coming contact with water, the tissue swells and closes the micropyle. Thus it prevents
the entry of water into the seeds.
(2) Prevents oxygen: Some seeds though they are capable of imbibing water, they are incapable
of taking in atmospheric air. In xanthium every fruit contains two seeds placed one above the
other. Curiously, the lower one germinates under normal conditions, but the upper one does not
until and unless, it is subjected to high oxygen pressure. This behavior has been attributed to the
presence of an inhibitor which will be oxidized only under higher concentration of
oxygen. Otherwise it inhibits the growth of the embryo.
(3) Prevents the growth of the embryo. Amaranthus and some other seeds belonging to this
category are capable of absorbing water and oxygen, with this the embryo is capable of growing, but
unfortunately it cannot break open the hard seed coat, thus the germination of seeds is prevented.
In this condition, the seeds may remain for months or years until the said seed coat gets cracked or
loosened.
In all the above mentioned cases the embryos are normal and do not posses any growth inhibiting
factors. But its growth is prevented by the presence of a hard seed coat which either prevents the
entry of water or oxygen or the seed coat does not crack and prevents the emergence of growing
embryo. The only mechanism by which such seeds can be induced to germinate is to break the seed
coat and make it weak so that the seeds can take up water and oxygen and facilitate the embryo to
emerge out of the seed coat. This can be achieved by a process called scarification which can be
done by following methods.
1. Mechanical Scarification: Shaking the seed with abrasives or nicking the seed coat with sharp
edged metals or chewing the seed coats without damaging the embryo makes the seed coat to crack
open. This greatly facilitates the emergence of the embryo out of the road coat.
2. Chemical Scarification: The hard seed coats can be loosened by strong acids or solvent
treatments where the hard coat is rendered soft. However the duration of treatment has to be
determined for every kind of seed, otherwise the treatment may cause damage to the young
embryos.
Many plants shed their seeds before the embroyos are fully mature. In such cases, germination fails
to occur till the embryos reach a maturity state. Maturation of such seeds can be achieved by
storing them under favorably conditions. But some seeds do not germinate even under such
favorable conditions. For that the storing has to be prolonged for some more time, only then the
seeds germinate. This behavior is often called after ripening effect. This behavior has been
explained as due to certain changes in its metabolic activities during storage. The maturation of
such seeds, noetheless can be augmented or accelerated by keeping layers of seeds alternating with
moist sand or moss at low temperatures. But the mechanism by which layering accelerates the
maturation of embryos is not known. But one probable explanation is that by keeping the seeds
under moist conditions many inhibitors are supposed to leach out.
Temperature Effect:
A large number of plants produce seeds which germinate under normal temperatures. But some do
not germinate if they are stored at room temperatures. They require chilling treatment for a period
of time. It is only then the seeds over come dormancy and germinate. Probably cold treatment
destroys the inhibitors present in the seed coat or in the embryo. This effect is almost like
vernalization, but the mechanism is different about which we dont know much.
Effect of Light:
Besides initiating many other photobiological processes, radiant energy has a profound influence on
seed dormancy and germination. Among the innumerable species of plant, some are in sensitive to
light radiations. Based on the above said property seeds have been classified into three kinds, i.e.
positive photoblastic types, negative photoblastic types and non photoblastic types.
In the case of positive photoblastic types when seeds are exposed to one or two cycles of intense
source of white light, they germinate. On the other hand, the negative photoblastic varieties do not
respond to white light treatment, but they germinate if they are maintained in complete dark
conditions. The non photoblastic types are insensitive to light and they germinate irrespective of the
presence or absence of light.
The effective spectrum of the visible light that induces dormancy or germination in light sensitive
seeds is red light and far red light. The seeds of Grand Rapids and Pepper grass germinate promptly
if they are exposed to red light. On the contrary, far red light inhibits their germination and seeds
remain dormant. However the red and far red light effects on seed germination can be
reversed. The table given below indicates that with the increase in the number of alternate cycles of
red and far red light treatments ending with red light, the percentage of seed germination enhances
significantly. But the treatment with far red as the last of the cycle, the percentage of inhibition is
more or less 90-93%.
The effectiveness of red light and far red light in inducing or inhibiting the germination further
suggests the involvement of phytochromes in this process. In fact, Ikuma and Thimann, way back,
demonstrated the role of phytochromes in inducing seed dormancy or germination. However, the
effectiveness of phytochromes depends upon other factors like water, temperature, pH, and age of
seeds, duration of light treatment and other chemicals present in seeds.
Phytochromes, as described elsewhere, exist in two forms which are interconvertible in the presence
of red and far red lights (PR and PfR).
The PfR produced under red light conditions undergoes reverse reaction in dark to produce PR form
of the pigment. However, in some cases it has been found that the PR form is converted to PfR form
by specific enzyme medicated reactions which require the input of energy. Furthermore, it is also
known that PfR requires another unknown factor called X to be in active state. (PfR-X).
The PfR-X form of pigment complex is believed to be very effective in breaking the dormancy in red
light requiring positive photoblastic seeds. The only difference between the positive photoblastic
seeds and negative photoblastic seeds is that the former requires higher concentration of PfR X
complex than the latter one; where as the negative photoblastic seeds require a minimum amount
of PfR complex to be highly effective. If the concentration of PfR is more in negative photoblastic
seeds their do not germinate. Correspondingly, the non-photoblastic varieties germinate without
light treatment, none the less PfR has a promotive effect. In spite of all these studies, the exact
mechanism by which the phytochrome pigments bring about dormancy or break the dormancy is
not clear.
Search for substances that induce dormancy in seeds resulted in the discovery of a host of
compounds like Coumerin, Para ascorbic acid, Hydrogen cyanide, Abscisic acid, etc. Most of them
have been isolated from the seed coats, endosperm pulp and the juice of fruits of embryos. In
those species which exhibits dormancy during the maturation of seeds and fruits some of the above
said compounds are actively synthesized and stored in different parts of seeds and fruits.
The above said substances are non toxic but induce dormancy. Until and unless the said compounds
are either destroyed or leached out, seeds remain dormant.
In recent years, the effect of ABA on seed drying and inducing dormancy in seeds has attracted a lot
of attention. In many fleshy fruits the ripening process is also initiated by ethylene. But ethylene
inturn can induce the synthesis of ABA. The abscissins thus produced bring about the induction of
few enzymes which not only facilitate fruit ripening but also induce changes in membrane
permeability and further they influences transcriptional activity as well as translational activity. Such
changes ultimately slow down most of the metabolic activities in the embryo. At the same time, the
water content of the embryonic cells is greatly reduced, resulting in the suspension of all metabolic
activities of the embryo, thus seeds become dormant. The degree of dormancy depends upon the
viable concentration of ABA stored in seeds.
Plants have unique properties in synthesizing compounds which can induce dormancy as well as
break the dormancy. Of course the synthesis of such compounds takes place at different
environmental conditions. Some of the compounds produced by plants are as effective as light and
temperature in breaking the seed dormancy. The compounds that are known to overcome
dormancy are gibberellins, cytokinins, ethylene, chlorohydrins, theourea, etc.
The action of Gibberellins is breaking seed dormancy is interesting because they are very effective
on seeds that require light treatment for germination. It is now known that both GA and ABA are
synthesized in the same plastids and their synthetic pathway starts from the common precursor
called Mevalonate. Furthermore, it can be demonstrated how the synthesis of GA and ABA is
correlated to red light and far red light mediated phytochrome activity
In red light requiring seeds on exposure to the red light more of PfR pigments accumulate. Once the
concentration of these pigments reaches a threshold value, they probably initiate or favor the
pathway of GA synthesis and also they facilitate the release of GA. The release of GA inturn
activates the respiratory activity and metabolic activities and activate certain genes (via transcription
and translation) resulting in the production of various components required for the active growth
and development of embryo. Thus GA overcomes the dormancy and induces germination. On the
contrary, short day or far red induced dormancy is due to the accumulation of more of PR form of
pigments. These pigments in fact favor the pathway of synthesis and release of ABA from
plastids. Then ABA brings about the inhibitory effect on cellular metabolism and imposes seed
dormancy. It is also speculated that each of these components produced in response to different
light treatments cant bring about feed back inhibition on each others pathway. The interplay of GA
and ABA in response to different light periods, either in breaking the dormancy or in imposing the
dormancy, is very interesting.