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Germination

Dormancy and life span of seeds

Dormancy has at least three functions: (1) immediate germination must be prevented even when circumstances are optimal so as to avoid exposure of the seedling to an unfavourable period (e.g., winter), which is sure to follow; (2) the unfavourable period has to be survived; and (3) the various dispersing agents must be given time to act. Accordingly, the wide variation in seed and diaspore longevity can be appreciated only by linking it with the various dispersal mechanisms employed as well as with the climate and its seasonal changes. Thus, the downy seeds of willows, blown up and down rivers in early summer with a chance of quick establishment on newly exposed sandbars, have a life span of only one week. Tropical rainforest trees frequently have seeds of low life expectancy also. Intermediate are seeds of sugarcane, tea, and coconut palm, among others, with life spans of up to a year. Mimosa glomerata seeds in the herbarium of the Muséum National d’Histoire Naturelle in Paris were found viable after 221 years. In general, viability is better retained in air of low moisture content. Some seeds, however, remain viable underwater—those of certain rush (Juncus) species and Sium cicutaefolium for at least 7 years. Salt water can be tolerated for years by the pebblelike but floating seeds of Guilandina bonduc, which in consequence possess an almost pantropical distribution. Seeds of the sacred lotus ( Nelumbo nucifera) found in a peat deposit in Manchuria and estimated by radioactive-carbon dating to be 1,400 ± 400 years old rapidly germinated (and subsequently produced flowering plants) when the seeds were filed to permit water entry. In 1967, seeds of the arctic tundra lupine ( Lupinus arcticus) found in a frozen lemming burrow with animal remains established to be at least 10,000 years old germinated within 48 hours when returned to favourable conditions. The problem of differential seed viability has been approached experimentally by various workers, one of whom buried 20 species of common Michigan weed seeds, mixed with sand, in inverted open-mouthed bottles for periodic inspection. After 80 years, 3 species still had viable seeds. See also soil seed bank.

Lack of dormancy

In some plants, the seeds are able to germinate as soon as they have matured on the plant, as demonstrated by papaya and by wheat, peas, and beans in a very rainy season. Certain mangrove species normally form foot-long embryos on the trees; these later drop down into the mud or sea water. Such cases, however, are exceptional. The lack of dormancy in cultivated species, contrasting with the situation in most wild plants, is undoubtedly the result of conscious selection by humans.

Immature embryos

In plants whose seeds ripen and are shed from the mother plant before the embryo has undergone much development beyond the fertilized egg stage (orchids, broomrapes, ginkgo, ash, winter aconite, and buttercups), there is an understandable delay of several weeks or months, even under optimal conditions, before the seedling emerges.

Role of the seed coat

There are at least three ways in which a hard testa may be responsible for seed dormancy: it may (1) prevent expansion of the embryo mechanically, (2) block the entrance of water, or (3) impede gas exchange so that the embryos lack oxygen. Resistance of the testa to water uptake is most widespread in the bean family, the seed coats of which, usually hard, smooth, or even glassy, may, in addition, possess a waxy covering. In some cases water entry is controlled by a small opening, the strophiolar cleft, which is provided with a corklike plug; only removal or loosening of the plug will permit water entry. Similar seeds not possessing a strophiolar cleft must depend on abrasion, which in nature may be brought about by microbial attack, passage through an animal, freezing and thawing, or mechanical means. In horticulture and agriculture, the coats of such seeds are deliberately damaged or weakened by humans ( scarification). In chemical scarification, seeds are dipped into strong sulfuric acid, organic solvents such as acetone or alcohol, or even boiling water. In mechanical scarification, they may be shaken with some abrasive material such as sand or be scratched with a knife.

Frequently seed coats are permeable to water yet block entrance of oxygen; this applies, for example, to the upper of the two seeds normally found in each burr of the cocklebur plant. The lower seed germinates readily under a favourable moisture and temperature regime, but the upper one fails to do so unless the seed coat is punctured or removed or the intact seed is placed under very high oxygen concentrations.

Seed – Seed – Germination: Dormancy has at least three functions: (1) immediate germination must be prevented even when circumstances are optimal so as to avoid exposure of the seedling to an unfavourable period (e.g., winter), which is sure to follow; (2) the unfavourable period has to be survived; and (3) the various dispersing agents must be given time to act. Accordingly, the wide variation in seed and diaspore longevity can be appreciated only by linking it with the various dispersal mechanisms employed as well as with the climate and its seasonal changes. Thus, the downy seeds of willows, blown up and down rivers in

Germination

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Germination, the sprouting of a seed, spore, or other reproductive body, usually after a period of dormancy. The absorption of water, the passage of time, chilling, warming, oxygen availability, and light exposure may all operate in initiating the process.

In the process of seed germination, water is absorbed by the embryo, which results in the rehydration and expansion of the cells. Shortly after the beginning of water uptake, or imbibition, the rate of respiration increases, and various metabolic processes, suspended or much reduced during dormancy, resume. These events are associated with structural changes in the organelles (membranous bodies concerned with metabolism), in the cells of the embryo.

Germination sometimes occurs early in the development process; the mangrove (Rhizophora) embryo develops within the ovule, pushing out a swollen rudimentary root through the still-attached flower. In peas and corn (maize) the cotyledons (seed leaves) remain underground (e.g., hypogeal germination), while in other species (beans, sunflowers, etc.) the hypocotyl (embryonic stem) grows several inches above the ground, carrying the cotyledons into the light, in which they become green and often leaflike (e.g., epigeal germination).

Seed dormancy

Dormancy is brief for some seeds—for example, those of certain short-lived annual plants. After dispersal and under appropriate environmental conditions, such as suitable temperature and access to water and oxygen, the seed germinates, and the embryo resumes growth.

The seeds of many species do not germinate immediately after exposure to conditions generally favourable for plant growth but require a “breaking” of dormancy, which may be associated with change in the seed coats or with the state of the embryo itself. Commonly, the embryo has no innate dormancy and will develop after the seed coat is removed or sufficiently damaged to allow water to enter. Germination in such cases depends upon rotting or abrasion of the seed coat in the gut of an animal or in the soil. Inhibitors of germination must be either leached away by water or the tissues containing them destroyed before germination can occur. Mechanical restriction of the growth of the embryo is common only in species that have thick, tough seed coats. Germination then depends upon weakening of the coat by abrasion or decomposition.

In many seeds the embryo cannot germinate even under suitable conditions until a certain period of time has lapsed. The time may be required for continued embryonic development in the seed or for some necessary finishing process—known as afterripening—the nature of which remains obscure.

The seeds of many plants that endure cold winters will not germinate unless they experience a period of low temperature, usually somewhat above freezing. Otherwise, germination fails or is much delayed, with the early growth of the seedling often abnormal. (This response of seeds to chilling has a parallel in the temperature control of dormancy in buds.) In some species, germination is promoted by exposure to light of appropriate wavelengths. In others, light inhibits germination. For the seeds of certain plants, germination is promoted by red light and inhibited by light of longer wavelength, in the “far red” range of the spectrum. The precise significance of this response is as yet unknown, but it may be a means of adjusting germination time to the season of the year or of detecting the depth of the seed in the soil. Light sensitivity and temperature requirements often interact, the light requirement being entirely lost at certain temperatures.

Seedling emergence

Active growth in the embryo, other than swelling resulting from imbibition, usually begins with the emergence of the primary root, known as the radicle, from the seed, although in some species (e.g., the coconut) the shoot, or plumule, emerges first. Early growth is dependent mainly upon cell expansion, but within a short time cell division begins in the radicle and young shoot, and thereafter growth and further organ formation (organogenesis) are based upon the usual combination of increase in cell number and enlargement of individual cells.

Until it becomes nutritionally self-supporting, the seedling depends upon reserves provided by the parent sporophyte. In angiosperms these reserves are found in the endosperm, in residual tissues of the ovule, or in the body of the embryo, usually in the cotyledons. In gymnosperms food materials are contained mainly in the female gametophyte. Since reserve materials are partly in insoluble form—as starch grains, protein granules, lipid droplets, and the like—much of the early metabolism of the seedling is concerned with mobilizing these materials and delivering, or translocating, the products to active areas. Reserves outside the embryo are digested by enzymes secreted by the embryo and, in some instances, also by special cells of the endosperm.

In some seeds (e.g., castor beans) absorption of nutrients from reserves is through the cotyledons, which later expand in the light to become the first organs active in photosynthesis. When the reserves are stored in the cotyledons themselves, these organs may shrink after germination and die or develop chlorophyll and become photosynthetic.

Environmental factors play an important part not only in determining the orientation of the seedling during its establishment as a rooted plant but also in controlling some aspects of its development. The response of the seedling to gravity is important. The radicle, which normally grows downward into the soil, is said to be positively geotropic. The young shoot, or plumule, is said to be negatively geotropic because it moves away from the soil; it rises by the extension of either the hypocotyl, the region between the radicle and the cotyledons, or the epicotyl, the segment above the level of the cotyledons. If the hypocotyl is extended, the cotyledons are carried out of the soil. If the epicotyl elongates, the cotyledons remain in the soil.

Light affects both the orientation of the seedling and its form. When a seed germinates below the soil surface, the plumule may emerge bent over, thus protecting its delicate tip, only to straighten out when exposed to light (the curvature is retained if the shoot emerges into darkness). Correspondingly, the young leaves of the plumule in such plants as the bean do not expand and become green except after exposure to light. These adaptative responses are known to be governed by reactions in which the light-sensitive pigment phytochrome plays a part. In most seedlings, the shoot shows a strong attraction to light, or a positive phototropism, which is most evident when the source of light is from one direction. Combined with the response to gravity, this positive phototropism maximizes the likelihood that the aerial parts of the plant will reach the environment most favourable for photosynthesis.

Germination, the sprouting of a seed, spore, or other reproductive body, usually after a period of dormancy. The absorption of water, the passage of time, chilling, warming, oxygen availability, and light exposure may all operate in initiating the process.