Seed dormancy
Encyclopedia
Seed dormancy is a condition of plant seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...

s that prevents germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

 when the seeds are under optimal environmental conditions for germination. Living, non dormant seeds germinate when soil temperatures and moisture conditions are suited for cellular processes and division; dormant seeds do not.

One important function of most seeds is delayed germination, which allows time for dispersal and prevents germination of all the seeds at same time. The staggering of germination safeguards some seeds and seedlings from suffering damage or death from short periods of bad weather or from transient herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

s; it also allows some seeds to germinate when competition from other plants for light and water might be less intense. Another form of delayed seed germination is seed quiescence, which is different than true seed dormancy and occurs when a seed fails to germinate because the external environmental conditions are too dry or warm or cold for germination. Many species of plants have seeds that delay germination for many months or years, and some seeds can remain in the soil seed bank for more than 50 years before germination. Some seeds have a very long viability period, and the oldest documented germinating seed was nearly 2000 years old based on radiocarbon dating
Radiocarbon dating
Radiocarbon dating is a radiometric dating method that uses the naturally occurring radioisotope carbon-14 to estimate the age of carbon-bearing materials up to about 58,000 to 62,000 years. Raw, i.e. uncalibrated, radiocarbon ages are usually reported in radiocarbon years "Before Present" ,...

.

Overview

True dormancy or innate dormancy is caused by conditions within the seed that prevent germination under normally ideal conditions. Often seed dormancy is divided into two major categories based on what part of the seed produces dormancy: exogenous and endogenous. There are three types of dormancy based on their mode of action: physical, physiological and morphological.

There have been a number of classification schemes developed to group different dormant seeds, but none have gained universal usage. Dormancy occurs because of a wide range of reasons that often overlap, producing conditions in which definitive categorization is not clear. Compounding this problem is that the same seed that is dormant for one reason at a given point may be dormant because of another reason at a later point. Some seeds fluctuate from periods of dormancy to non dormancy, and despite the fact that a dormant seed appears to be static or inert, in reality they are still receiving and responding to environmental cues.

Exogenous dormancy

Exogenous dormancy is caused by conditions outside the embryo and is often broken down into three subgroups:

Physical dormancy

Which occurs when seeds are impermeable
Permeability (fluid)
Permeability in fluid mechanics and the earth sciences is a measure of the ability of a porous material to allow fluids to pass through it.- Units :...

 to water or the exchange of gases. Legumes are typical examples of physically dormant seeds; they have low moisture content and are prevented from imbibing water by the seed coat. Chipping or cracking of the seed coat or any other coverings allows water intake. Impermeability is often caused by an outer cell layer which is composed of macrosclereid cells or the outer layer is composed of a mucilaginous cell layer. The third cause of seed coat impermeability is a hardened endocarp. Seed coats that are impermeable to water and gases form during the last stages of seed development.

Mechanical dormancy

Mechanical dormancy occurs when seed coats or other coverings are too hard to allow the embryo to expand during germination. In the past this mechanism of dormancy was ascribed to a number of species that have been found to have endogenous factors for their dormancy instead. These endogenous facts include physiologically dormancy cased by low embryo growth potential

Chemical dormancy

Includes growth regulators etc., that are present in the coverings around the embryo. They may be leached out of the tissues by washing or soaking the seed, or deactivated by other means. Other chemicals that prevent germination are washed out of the seeds by rainwater or snow melt.

Endogenous dormancy

Endogenous dormancy is caused by conditions within the embryo itself, and it is also often broken down into three subgroups: physiological dormancy, morphological dormancy and combined dormancy, each of these groups may also have subgroups.

Physiological dormancy

Physiological dormancy prevents embryo growth and seed germination until chemical changes occur. These chemicals include inhibitors that often retard embryo growth to the point where it is not strong enough to break through the seed coat or other tissues. Physiological dormancy is indicated when an increase in germination rate occurs after an application of gibberellic acid
Gibberellic acid
Gibberellic acid Gibberellic acid Gibberellic acid (also called Gibberellin A3, GA, and (GA3) is a hormone found in plants. Its chemical formula is C19H22O6. When purified, it is a white-to-pale-yellow solid....

 (GA3) or after Dry after-ripening or dry storage. It is also indicated when dormant seed embryos are excised and produce healthy seedlings: or when up to 3 months of cold (0-10°C) or warm (=15°C) stratification increases germination: or when dry after-ripening shortens the cold stratification period required. In some seeds physiological dormancy is indicated when scarification increases germination.

Physiological dormancy is broken when inhibiting chemicals are broken down or are no longer produced by the seed; often by a period of cool moist conditions, normally below (+4C) 39F, or in the case of many species in Ranunculaceae
Ranunculaceae
Ranunculaceae are a family of about 1700 species of flowering plants in about 60 genera, distributed worldwide....

and a few others,(-5C) 24F. Abscisic acid
Abscisic acid
Abscisic acid , also known as abscisin II and dormin, is a plant hormone. ABA functions in many plant developmental processes, including bud dormancy. It is degraded by the enzyme -abscisic acid 8'-hydroxylase.-Function:...

 is usually the growth inhibitor in seeds and its production can be affected by light. Some plants like Peony
Peony
Peony or paeony is a name for plants in the genus Paeonia, the only genus in the flowering plant family Paeoniaceae. They are native to Asia, southern Europe and western North America...

 species have multiple types of physiological dormancy, one affects radicle (root) growth while the other affects plumule (shoot) growth. Seeds with physiological dormancy most often do not germinate even after the seed coat or other structures that interfere with embryo growth are removed. Conditions that affect physiological dormancy of seeds include:
  • 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 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.
  • 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
    Light
    Light or visible light is electromagnetic radiation that is visible to the human eye, and is responsible for the sense of sight. Visible light has wavelength in a range from about 380 nanometres to about 740 nm, with a frequency range of about 405 THz to 790 THz...

     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
    Cocklebur
    Cockleburs are a genus of flowering plants in the family Asteraceae, native to the Americas and eastern Asia.-Growth:They are coarse, herbaceous annual plants growing to 19.69-47.24 in tall. The leaves are spirally arranged, with a deeply toothed margin. Some species, notably X...

     and amaranth
    Amaranth
    Amaranthus, collectively known as amaranth, is a cosmopolitan genus of herbs. Approximately 60 species are recognized, with inflorescences and foliage ranging from purple and red to gold...

     germinate only at high temperatures (30C or 86F) many plants that have seed that germinate in early to mid summer have thermodormancy 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 thermodormancy requirements disappear as the seed ages or dries.

Seeds are classified as having deep physiological dormancy under these conditions: applications of GA3 does not increase germination; or when excised embryos produce abnormal seedlings; or when seeds require more than 3 months of cold stratification to germinate.

Morphological dormancy

Embryo underdeveloped or undifferentiated. Some seeds have fully differentiated embryos that need to grow more before seed germination, or the embryos are not differentiated into different tissues at the time of fruit ripening.
  • Immature embryos - some plants release their seeds before the tissues of the embryos have fully differentiated, and the seeds ripen after they take in water while on the ground, germination can be delayed from a few weeks to a few months.

Combined dormancy

Seeds have both morphological and physiological dormancy.
  • Morpho-physiological or morphophysiological dormancy occurs when seeds with underdeveloped embryos, also have physiological components to dormancy. These seeds therefore require dormancy-breaking treatments as well as a period of time to develop fully grown embryos.
  • Intermediate simple -
  • Deep simple -
  • Deep simple epicotyl -
  • Deep simple double -
  • Intermediate complex -
  • Deep complex -

Combinational dormancy

Combinational dormancy occurs in some seeds, where dormancy is caused by both exogenous (physical) and
endogenous (physiological) conditions. some Iris
Iris (plant)
Iris is a genus of 260-300species of flowering plants with showy flowers. It takes its name from the Greek word for a rainbow, referring to the wide variety of flower colors found among the many species...

species have both hard impermeable seeds coats and physiological dormancy.

Secondary dormancy

Secondary dormancy occurs in some non-dormant and post dormant seeds that are exposed to conditions that are not favorable for germination, like high temperatures. It is caused by conditions that occur after the seed has been dispersed. The mechanisms of secondary dormancy are not yet fully understood but might involve the loss of sensitivity in receptors in the plasma membrane.

Not all seeds undergo a period of dormancy, many species of plants release their seeds late in the year when the soil temperature is too low for germination or when the environment is dry. If these seeds are collected and sown in an environment that is warm enough, and/or moist enough, they will germinate. Under natural conditions non dormant seeds released late in the growing season wait until spring when the soil temperature rises or in the case of seeds dispersed during dry periods until it rains and there is enough soil moisture.

Seeds that do not germinate because they have fleshy fruits that retard germination are quiescent, not dormant.

Many garden plants have seeds that will germinate readily as soon as they have water and are warm enough, though their wild ancestors had dormancy. These cultivated plants lack seed dormancy because of generations of selective pressure by plant breeders and gardeners that grew and kept plants that lacked dormancy.

Seeds of some mangrove
Mangrove
Mangroves are various kinds of trees up to medium height and shrubs that grow in saline coastal sediment habitats in the tropics and subtropics – mainly between latitudes N and S...

s are viviparous and begin to germinate while still attached to the parent; they produce a large, heavy root, which allows the seed to penetrate into the ground when it falls.
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