Seed - Seed Functions - Seed Dormancy

Seed Dormancy

Seed dormancy has two main functions: the first is synchronizing germination with the optimal conditions for survival of the resulting seedling; the second is spreading germination of a batch of seeds over time so a catastrophe after germination (e.g. late frosts, drought, herbivory) does not result in the death of all offspring of a plant (bet-hedging). Seed dormancy is defined as a seed failing to germinate under environmental conditions optimal for germination, normally when the environment is at a suitable temperature with proper soil moisture. This true dormancy or innate dormancy is therefore caused by conditions within the seed that prevent germination. Thus dormancy is a state of the seed, not of the environment. Induced dormancy, enforced dormancy or seed quiescence occurs when a seed fails to germinate because the external environmental conditions are inappropriate for germination, mostly in response to conditions being too dark or light, too cold or hot, or too dry.

Seed dormancy is not the same as seed persistence in the soil or on the plant, though even in scientific publications dormancy and persistence are often confused or used as synonyms.

Often, seed dormancy is divided into four major categories: exogenous; endogenous; combinational; and secondary. A more recent system distinguishes five classes: 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 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, but little evidence exists to support this claim.

Endogenous dormancy is caused by conditions within the embryo itself, including:

• In morphological dormancy, 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 (underdeveloped) 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 includes seeds with underdeveloped embryos, and also 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, such as Asarum or Trillium species, have multiple types of dormancy, one affects radicle (root) growth, while the other affects plumule (shoot) growth. The terms "double dormancy" and "two-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 the embryo, due to physiological causes, cannot 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, in some plants, including a number of grasses and those from seasonally arid regions, is needed before they will germinate. The seeds are released, but need to have a 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.

• 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 nondormant 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.

The following types of seed dormancy do not involve seed dormancy, strictly speaking, as lack of germination is prevented by the environment, not by characteristics of the seed itself (see Germination):

• Photodormancy 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.
• Thermodormancy is seed sensitivity to heat or cold. Some seeds, including cocklebur and amaranth, germinate only at high temperatures (30°C or 86°F); many plants that have seeds that germinate in early to midsummer have thermodormancy, so germinate only when the soil temperature is warm. Other seeds need cool soils to germinate, while others, such as celery, are inhibited when soil temperatures are too warm. Often, thermodormancy 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 plant seeds 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 it. 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.