Seed Dormancy
What is seed dormancy and why it is important to know
Plant hormones affect seed germination and dormancy by affecting different parts of the seed.
Embryo dormancy is characterised by a high ABA/GA ratio, whereas the seed has a high ABA sensitivity and low GA sensitivity. To release the seed from this type of dormancy and initiate seed germination, an alteration in hormone biosynthesis and degradation towards a low ABA/GA ratio, along with a decrease in ABA sensitivity and an increase in GA sensitivity needs to occur.
ABA controls embryo dormancy, and GA embryo germination. Seed coat dormancy involves the mechanical restriction of the seed coat, this along with a low embryo growth potential, effectively produces seed dormancy. GA releases this dormancy by increasing the embryo growth potential, and/or weakening the seed coat so the radical of the seedling can break through the seed coat. Different types of seed coats can be made up of living or dead cells and both types can be influenced by hormones; those composed of living cells are acted upon after seed formation while the seed coats composed of dead cells can be influenced by hormones during the formation of the seed coat. ABA affects testa or seed coat growth characteristics, including thickness, and effects the GA-mediated embryo growth potential. These conditions and effects occur during the formation of the seed, often in response to environmental conditions.
Hormones also mediate endosperm dormancy: Endosperm in most seeds is composed of living tissue that can actively respond to hormones generated by the embryo. The endosperm often acts as a barrier to seed germination, playing a part in seed coat dormancy or in the germination process. Living cells respond to and also affect the ABA/GA ratio, and mediate cellular sensitivity; GA thus increases the embryo growth potential and can promote endosperm weakening. GA also affects both ABA-independent and ABA-inhibiting processes within the endosperm.
Often seed dormancy is divided into four major categories: exogenous; endogenous; combinational; and secondary. A more recent system distinguishes five classes of dormancy: morphological, physiological, morphophysiological, physical and combinational dormancy.
Exogenous dormancy is caused by conditions outside the embryo including:
Physical dormancy or hard seed coats occurs when seeds are impermeable to water. At dormancy break a specialized structure, the ‘water gap’, is disrupted in response to environmental cues, especially temperature, so that water can enter the seed and germination can occur. Plant families where physical dormancy occurs include Anacardiaceae, Cannaceae, Convulvulaceae, Fabaceae and Malvaceae.
Chemical dormancy considers species that lack physiological dormancy, but where a chemical prevents germination. This chemical can be leached out of the seed by rainwater or snow melt or be deactivated somehow. Leaching of chemical inhibitors from the seed by rain water is often cited as an important cause of dormancy release in seeds of desert plants, however little evidence exists to support this claim.
Endogenous dormancy is caused by conditions within the embryo itself, including:
Morphological dormancy where germination is prevented due to morphological characteristics of the embryo. In some species the embryo is just a mass of cells when seeds are dispersed, it is not differentiated. Before germination can take place both differentiation and growth of the embryo have to occur. In other species the embryo is differentiated but not fully grown at dispersal and embryo growth up to a species specific length is required before germination can occur. Examples of plant families where morphological dormancy occurs are Apiaceae, Cycadaceae, Liliaceae, Magnoliaceae and Ranunculaceae.
Morphophysiological dormancy seeds with underdeveloped embryos and which in addition have physiological components to dormancy. These seeds therefore require a dormancy-breaking treatments as well as a period of time to develop fully grown embryos. Plant families where morphophysiological dormancy occurs include Apiaceae, Aquifoliaceae, Liliaceae, Magnoliaceae, Papaveraceae and Ranunculaceae. Some plants with morphophysiological dormancy like Asarum or Trillium species have multiple types of dormancy, one affects radicle growth while the other affects plumule or shoot growth. The terms double dormancy and 2-year seeds are used for species whose seeds need two years to complete germination or at least two winters and one summer. Dormancy of the radicle (seedling root)is broken during the first winter after dispersal while dormancy of the shoot bud is broken during the second winter.
Physiological dormancy means that the embryo can, due to physiological causes, not generate enough power to break through the seed coat, endosperm or other covering structures. Dormancy is typically broken at cool wet, warm wet or warm dry conditions. Abscisic acid is usually the growth inhibitor in seeds and its production can be affected by light.
Drying; some plants including a number of grasses and those from seasonally arid regions need a period of drying before they will germinate, the seeds are released but need to have lower moisture content before germination can begin. If the seeds remain moist after dispersal, germination can be delayed for many months or even years. Many herbaceous plants from temperate climate zones have physiological dormancy that disappears with drying of the seeds. Other species will germinate after dispersal only under very narrow temperature ranges, but as the seeds dry they are able to germinate over a wider temperature range.
Combinational dormancy In seeds with combinational dormancy the seed or fruit coat is impermeable to water and the embryo has physiological dormancy. Depending on the species physical dormancy can be broken before or after physiological dormancy is broken.
Secondary dormancy is caused by conditions after the seed has been dispersed and occurs in some seeds when non-dormant seed is exposed to conditions that are not favorable to germination, very often high temperatures. The mechanisms of secondary dormancy are not yet fully understood but might involve the loss of sensitivity in receptors in the plasma membrane.
Additional dormancy strategies
The following types of seed dormancy do not involve seed dormancy strictly spoken as lack of germination is prevented by the environment not by characteristics of the seed itself:
Photo-dormancy or light sensitivity affects germination of some seeds. These photoblastic seeds need a period of darkness or light to germinate. In species with thin seed coats, light may be able to penetrate into the dormant embryo. The presence of light or the absence of light may trigger the germination process, inhibiting germination in some seeds buried too deeply or in others not buried in the soil.
Thermo-dormancy is seed sensitivity to heat or cold. Some seeds including cocklebur and amaranth germinate only at high temperatures (30C or 86F) many plants that have seed that germinate in early to mid summer have thermo-dormancy and germinate only when the soil temperature is warm. Other seeds need cool soils to germinate, while others like celery are inhibited when soil temperatures are too warm. Often thermo-dormancy requirements disappear as the seed ages or dries.
Not all seeds undergo a period of dormancy. Seeds of some mangroves are viviparous, they begin to germinate while still attached to the parent. The large, heavy root allows the seed to penetrate into the ground when it falls. Many garden plants have seeds that will germinate readily as soon as they have water and are warm enough, though their wild ancestors may have had dormancy, these cultivated plants lack seed dormancy. After many generations of selective pressure by plant breeders and gardeners dormancy has been selected out.
For annuals, seeds are a way for the species to survive dry or cold seasons. Ephemeral plants are usually annuals that can go from seed to seed in as few as six weeks.