A seed is an embryonic plant enclosed in a protective outer covering.

The formation of the seed

is part of the process of reproduction in seed plants, the spermatophytes, including the
gymnosperm and angiosperm plants.

Seeds are the product of the ripened ovule, after fertilization by pollen and some growth within
the mother plant. The embryo is developed from the zygote and the seed coat from the
integuments of the ovule.

Seeds have been an important development in the reproduction and success of gymnosperm and
angiosperm plants, relative to more primitive plants such as ferns, mosses and liverworts, which
do not have seeds and use water-dependent means to propagate themselves. Seed plants now
dominate biological niches on land, from forests to grasslands both in hot and cold climates.

The term "seed" also has a general meaning that antedates the above – anything that can be
sown, e.g. "seed" potatoes, "seeds" of corn or sunflower "seeds". In the case of sunflower and
corn "seeds", what is sown is the seed enclosed in a shell or husk, whereas the potato is a tuber.

Many structures commonly referred to as "seeds" are actually dry fruits. Plants producing berries
are called baccate. Sunflower seeds are sometimes sold commercially while still enclosed within
the hard wall of the fruit, which must be split open to reach the seed. Different groups of plants
have other modifications, the so-called stone fruits (such as the peach) have a hardened fruit
layer (the endocarp) fused to and surrounding the actual seed. Nuts are the one-seeded, hard-
shelled fruit of some plants with an indehiscent seed, such as an acorn or hazelnut.

Seed production[edit]
Seeds are produced in several related groups of plants, and their manner of production
distinguishes the angiosperms ("enclosed seeds") from the gymnosperms ("naked seeds").
Angiosperm seeds are produced in a hard or fleshy structure called a fruit that encloses the seeds
for protection in order to secure healthy growth. Some fruits have layers of both hard and fleshy
material. In gymnosperms, no special structure develops to enclose the seeds, which begin their
development "naked" on the bracts of cones. However, the seeds do become covered by the cone
scales as they develop in some species of conifer.

Seed production in natural plant populations varies widely from year to year in response to
weather variables, insects and diseases, and internal cycles within the plants themselves. Over a
20-year period, for example, forests composed of loblolly pine and shortleaf pine produced from
0 to nearly 5 million sound pine seeds per hectare.[1] Over this period, there were six bumper, five
poor, and nine good seed crops, when evaluated for production of adequate seedlings for natural
forest reproduction.

Development[edit]
Stages of seed development:
I Zygote IV Heart
II Proembryo V Torpedo
III Globular VI Mature Embryo
Key: 1. Endosperm 2. Zygote 3. Embryo 4. Suspensor 5. Cotyledons 6. Shoot Apical Meristem 7.
Root Apical Meristem 8. Radicle 9. Hypocotyl 10. Epicotyl 11. Seed Coat

Angiosperm (flowering plants) seeds consist of three genetically distinct constituents: (1) the
embryo formed from the zygote, (2) the endosperm, which is normally triploid, (3) the seed coat
from tissue derived from the maternal tissue of the ovule. In angiosperms, the process of seed
development begins with double fertilization, which involves the fusion of two male gametes
with the egg cell and the central cell to form the primary endosperm and the zygote. Right after
fertilization, the zygote is mostly inactive, but the primary endosperm divides rapidly to form the
endosperm tissue. This tissue becomes the food the young plant will consume until the roots
have developed after germination.

Ovule[edit]

Main article: Ovule

Plant ovules: Gymnosperm ovule on left, angiosperm ovule (inside ovary) on right

After fertilization the ovules develop into the seeds. The ovule consists of a number of
components:
  The funicle (funiculus, funiculi) or seed stalk which attaches the ovule to the placenta and
hence ovary or fruit wall, at the pericarp.
 The nucellus, the remnant of the megasporangium and main region of the ovule where
the megagametophyte develops.
 The micropyle, a small pore or opening in the apex of the integument of the ovule where
the pollen tube usually enters during the process of fertilization.
 The chalaza, the base of the ovule opposite the micropyle, where integument and
nucellus are joined together.[2]

The shape of the ovules as they develop often affects the final shape of the seeds. Plants
generally produce ovules of four shapes: the most common shape is called anatropous, with a
curved shape. Orthotropous ovules are straight with all the parts of the ovule lined up in a long
row producing an uncurved seed. Campylotropous ovules have a curved megagametophyte
often giving the seed a tight "C" shape. The last ovule shape is called amphitropous, where the
ovule is partly inverted and turned back 90 degrees on its stalk (the funicle or funiculus).

In the majority of flowering plants, the zygote's first division is transversely oriented in regards
to the long axis, and this establishes the polarity of the embryo. The upper or chalazal pole
becomes the main area of growth of the embryo, while the lower or micropylar pole produces the
stalk-like suspensor that attaches to the micropyle. The suspensor absorbs and manufactures
nutrients from the endosperm that are used during the embryo's growth.[3]

Embryo[edit]

The inside of a Ginkgo seed, showing a well-developed embryo, nutritive tissue

(megagametophyte), and a bit of the surrounding seed coat

The main components of the embryo are:

 The cotyledons, the seed leaves, attached to the embryonic axis. There may be one
(Monocotyledons), or two (Dicotyledons). The cotyledons are also the source of nutrients
in the non-endospermic dicotyledons, in which case they replace the endosperm, and are
thick and leathery. In endospermic seeds the cotyledons are thin and papery.
Dicotyledons have the point of attachment opposite one another on the axis.
 The epicotyl, the embryonic axis above the point of attachment of the cotyledon(s).
 The plumule, the tip of the epicotyl, and has a feathery appearance due to the presence of
young leaf primordia at the apex, and will become the shoot upon germination.
  The hypocotyl, the embryonic axis below the point of attachment of the cotyledon(s),
connecting the epicotyl and the radicle, being the stem-root transition zone.
 The radicle, the basal tip of the hypocotyl, grows into the primary root.

Monocotyledonous plants have two additional structures in the form of sheaths. The plumule is
covered with a coleoptile that forms the first leaf while the radicle is covered with a coleorhiza
that connects to the primary root and adventitious roots form from the sides. Here the hypocotyl
is a rudimentary axis between radicle and plumule. The seeds of corn are constructed with these
structures; pericarp, scutellum (single large cotyledon) that absorbs nutrients from the
endosperm, plumule, radicle, coleoptile and coleorhiza – these last two structures are sheath-like
and enclose the plumule and radicle, acting as a protective covering.

Seed coat[edit]
The maturing ovule undergoes marked changes in the integuments, generally a reduction and
disorganization but occasionally a thickening. The seed coat forms from the two integuments or
outer layers of cells of the ovule, which derive from tissue from the mother plant, the inner
integument forms the segment and the outer forms the testa. (The seed coats of some
monocotyledon plants, such as the grasses, are not distinct structures, but are fused with the fruit
wall to form a pericardia.) The testae of both monocots and dicots are often marked with patterns
and textured markings, or have wings or tufts of hair. When the seed coat forms from only one
layer, it is also called the testa, though not all such testate are homologous from one species to
the next. The funiculars abscesses (detaches at fixed point – abscission zone), the scar forming
an oval depression, the hilum. Anatropous ovules have a portion of the funiculars that is adnate
(fused to the seed coat), and which forms a longitudinal ridge, or raphe, just above the hilum. In
bitegmic ovules (e.g. Symposium described here) both inner and outer integuments contribute to
the seed coat formation. With continuing maturation the cells enlarge in the outer integument.
While the inner epidermis may remain a single layer, it may also divide to produce two to three
layers and accumulates starch, and is referred to as the colourless layer. By contrast the outer
epidermis becomes tanniferous. The inner integument may consist of eight to fifteen layers.
(Kozlowski 1972)

As the cells enlarge, and starch is deposited in the outer layers of the pigmented zone below the
outer epidermis, this zone begins to lignify, while the cells of the outer epidermis enlarge radially
and their walls thicken, with nucleus and cytoplasm compressed into the outer layer. these cells
which are broader on their inner surface are called palisade cells. In the inner epidermis the cells
also enlarge radially with plate like thickening of the walls. The mature inner integument has a
palisade layer, a pigmented zone with 15–20 layers, while the innermost layer is known as the
fringe layer. (Kozlowski 1972)

In gymnosperms, which do not form ovaries, the ovules and hence the seeds are exposed. This is
the basis for their nomenclature – naked seeded plants. Two sperm cells transferred from the
pollen do not develop the seed by double fertilization, but one sperm nucleus unites with the egg
nucleus and the other sperm is not used. [4] Sometimes each sperm fertilizes an egg cell and one
zygote is then aborted or absorbed during early development.[5] The seed is composed of the
embryo (the result of fertilization) and tissue from the mother plant, which also form a cone
around the seed in coniferous plants such as pine and spruce.

Shape and appearance[edit]

A large number of terms are used to describe seed shapes, many of which are largely self-
explanatory such as Bean-shaped (reniform) – resembling a kidney, with lobed ends on either
side of the hilum, Square or Oblong – angular with all sides more or less equal or longer than
wide, Triangular – three sided, broadest below middle, Elliptic or Ovate or Obovate – rounded at
both ends, or egg shaped (ovate or obovate, broader at one end), being rounded but either
symmetrical about the middle or broader below the middle or broader above the middle.[6]

Other less obvious terms include discoid (resembling a disc or plate, having both thickness and
parallel faces and with a rounded margin), ellipsoid, globose (spherical), or subglobose
(Inflated, but less than spherical), lenticular, oblong, ovoid, reniform and sectoroid. Striate
seeds are striped with parallel, longitudinal lines or ridges. The commonest colours are brown
and black, other colours are infrequent. The surface varies from highly polished to considerably
roughened. The surface may have a variety of appendages (see Seed coat). A seed coat with the
consistency of cork is referred to as suberose. Other terms include crustaceous (hard, thin or
brittle).

Structure[edit]

The parts of an avocado seed (a dicot), showing the seed coat and embryo

Size and seed set[edit]

A collection of various vegetable and herb seeds

Seeds are very diverse in size. The dust-like orchid seeds are the smallest, with about one million
seeds per gram; they are often embryonic seeds with immature embryos and no significant
energy reserves. Orchids and a few other groups of plants are mycoheterotrophs which depend
on mycorrhizal fungi for nutrition during germination and the early growth of the seedling. Some
terrestrial orchid seedlings, in fact, spend the first few years of their lives deriving energy from
the fungi and do not produce green leaves.[14] At over 20 kg, the largest seed is the coco de mer.
Plants that produce smaller seeds can generate many more seeds per flower, while plants with
larger seeds invest more resources into those seeds and normally produce fewer seeds. Small
seeds are quicker to ripen and can be dispersed sooner, so fall blooming plants often have small
seeds. Many annual plants produce great quantities of smaller seeds; this helps to ensure at least
a few will end in a favorable place for growth. Herbaceous perennials and woody plants often
have larger seeds; they can produce seeds over many years, and larger seeds have more energy
reserves for germination and seedling growth and produce larger, more established seedlings
after germination.[15][16]

Functions[edit]
Seeds serve several functions for the plants that produce them. Key among these functions are
nourishment of the embryo, dispersal to a new location, and dormancy during unfavorable
conditions. Seeds fundamentally are means of reproduction, and most seeds are the product of
sexual reproduction which produces a remixing of genetic material and phenotype variability on
which natural selection acts.

Embryo nourishment[edit]

Seeds protect and nourish the embryo or young plant. They usually give a seedling a faster start
than a sporeling from a spore, because of the larger food reserves in the seed and the
multicellularity of the enclosed embryo.

Dispersal[edit]

Main article: Seed dispersal

Unlike animals, plants are limited in their ability to seek out favorable conditions for life and
growth. As a result, plants have evolved many ways to disperse their offspring by dispersing
their seeds (see also vegetative reproduction). A seed must somehow "arrive" at a location and be
there at a time favorable for germination and growth. When the fruits open and release their
seeds in a regular way, it is called dehiscent, which is often distinctive for related groups of
plants; these fruits include capsules, follicles, legumes, silicles and siliques. When fruits do not
open and release their seeds in a regular fashion, they are called indehiscent, which include the
fruits achenes, caryopsis, nuts, samaras, and utricles.[17]
By wind (anemochory)[edit]

Dandelion seeds are contained within achenes, which can be carried long distances by the wind.

The seed pod of milkweed (Asclepias syriaca)

 Some seeds (e.g., pine) have a wing that aids in wind dispersal.
 The dustlike seeds of orchids are carried efficiently by the wind.
 Some seeds (e.g. milkweed, poplar) have hairs that aid in wind dispersal.[18]

Other seeds are enclosed in fruit structures that aid wind dispersal in similar ways:

 Dandelion achenes have hairs.

 Maple samaras have two wings.

By water (hydrochory)[edit]

 Some plants, such as Mucuna and Dioclea, produce buoyant seeds termed sea-beans or drift
seeds because they float in rivers to the oceans and wash up on beaches.[19]
By animals (zoochory)[edit]

 Seeds (burrs) with barbs or hooks (e.g. acaena, burdock, dock) which attach to animal fur or
feathers, and then drop off later.
 Seeds with a fleshy covering (e.g. apple, cherry, juniper) are eaten by animals (birds, mammals,
reptiles, fish) which then disperse these seeds in their droppings.
 Seeds (nuts) are attractive long-term storable food resources for animals (e.g. acorns, hazelnut,
walnut); the seeds are stored some distance from the parent plant, and some escape being
eaten if the animal forgets them.

Myrmecochory is the dispersal of seeds by ants. Foraging ants disperse seeds which have
appendages called elaiosomes[20] (e.g. bloodroot, trilliums, acacias, and many species of
Proteaceae). Elaiosomes are soft, fleshy structures that contain nutrients for animals that eat
them. The ants carry such seeds back to their nest, where the elaiosomes are eaten. The
remainder of the seed, which is hard and inedible to the ants, then germinates either within the
nest or at a removal site where the seed has been discarded by the ants.[21] This dispersal
relationship is an example of mutualism, since the plants depend upon the ants to disperse seeds,
while the ants depend upon the plants seeds for food. As a result, a drop in numbers of one
partner can reduce success of the other. In South Africa, the Argentine ant (Linepithema humile)
has invaded and displaced native species of ants. Unlike the native ant species, Argentine ants do
not collect the seeds of Mimetes cucullatus or eat the elaiosomes. In areas where these ants have
invaded, the numbers of Mimetes seedlings have dropped.[22]

Dormancy[edit]

Main article: 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 (e.g. late frosts, drought, herbivory) does not result in the
death of all offspring of a plant (bet-hedging).[23] 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.[24] 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.[25]

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.[26]
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.[27]
 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.[28] 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.[29]

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.[30][31]
 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.[30] 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.[30]
 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.
o 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.[32]
 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.[31]
  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.[33]

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.[34]

Persistence and seed banks[edit]

Further information: Seed dormancy

Germination[edit]
Main articles: Seedling and Germination
Germinating sunflower seedlings

Seed germination is a process by which a seed embryo develops into a seedling. It involves the
reactivation of the metabolic pathways that lead to growth and the emergence of the radicle or
seed root and plumule or shoot. The emergence of the seedling above the soil surface is the next
phase of the plant's growth and is called seedling establishment.[35]

Three fundamental conditions must exist before germination can occur. (1) The embryo must be
alive, called seed viability. (2) Any dormancy requirements that prevent germination must be
overcome. (3) The proper environmental conditions must exist for germination.

Seed viability is the ability of the embryo to germinate and is affected by a number of different
conditions. Some plants do not produce seeds that have functional complete embryos, or the seed
may have no embryo at all, often called empty seeds. Predators and pathogens can damage or kill
the seed while it is still in the fruit or after it is dispersed. Environmental conditions like flooding
or heat can kill the seed before or during germination. The age of the seed affects its health and
germination ability: since the seed has a living embryo, over time cells die and cannot be
replaced. Some seeds can live for a long time before germination, while others can only survive
for a short period after dispersal before they die.

Seed vigor is a measure of the quality of seed, and involves the viability of the seed, the
germination percentage, germination rate and the strength of the seedlings produced.[36]

The germination percentage is simply the proportion of seeds that germinate from all seeds
subject to the right conditions for growth. The germination rate is the length of time it takes for
the seeds to germinate. Germination percentages and rates are affected by seed viability,
dormancy and environmental effects that impact on the seed and seedling. In agriculture and
horticulture quality seeds have high viability, measured by germination percentage plus the rate
of germination. This is given as a percent of germination over a certain amount of time, 90%
germination in 20 days, for example. 'Dormancy' is covered above; many plants produce seeds
with varying degrees of dormancy, and different seeds from the same fruit can have different
degrees of dormancy.[37] It's possible to have seeds with no dormancy if they are dispersed right
away and do not dry (if the seeds dry they go into physiological dormancy). There is great
variation amongst plants and a dormant seed is still a viable seed even though the germination
rate might be very low.
Environmental conditions affecting seed germination include; water, oxygen, temperature and
light.

Three distinct phases of seed germination occur: water imbibition; lag phase; and radicle
emergence.

In order for the seed coat to split, the embryo must imbibe (soak up water), which causes it to
swell, splitting the seed coat. However, the nature of the seed coat determines how rapidly water
can penetrate and subsequently initiate germination. The rate of imbibition is dependent on the
permeability of the seed coat, amount of water in the environment and the area of contact the
seed has to the source of water. For some seeds, imbibing too much water too quickly can kill the
seed. For some seeds, once water is imbibed the germination process cannot be stopped, and
drying then becomes fatal. Other seeds can imbibe and lose water a few times without causing ill
effects, but drying can cause secondary dormancy.

Repair of DNA damage[edit]

During seed dormancy, often associated with unpredictable and stressful environments, DNA
damage accumulates as the seeds age.[38][39][40] In rye seeds, the reduction of DNA integrity due to
damage is associated with loss of seed viability during storage.[38] Upon germination, seeds of
Vicia faba undergo DNA repair.[39] A plant DNA ligase that is involved in repair of single- and
double-strand breaks during seed germination is an important determinant of seed longevity.[41]
Also, in Arabidopsis seeds, the activities of the DNA repair enzymes Poly ADP ribose
polymerases (PARP) are likely needed for successful germination.[42] Thus DNA damages that
accumulate during dormancy appear to be a problem for seed survival, and the enzymatic repair
of DNA damages during germination appears to be important for seed viability.

Inducing germination[edit]

A number of different strategies are used by gardeners and horticulturists to break seed
dormancy.

Scarification allows water and gases to penetrate into the seed; it includes methods to physically
break the hard seed coats or soften them by chemicals, such as soaking in hot water or poking
holes in the seed with a pin or rubbing them on sandpaper or cracking with a press or hammer.
Sometimes fruits are harvested while the seeds are still immature and the seed coat is not fully
developed and sown right away before the seed coat become impermeable. Under natural
conditions, seed coats are worn down by rodents chewing on the seed, the seeds rubbing against
rocks (seeds are moved by the wind or water currents), by undergoing freezing and thawing of
surface water, or passing through an animal's digestive tract. In the latter case, the seed coat
protects the seed from digestion, while often weakening the seed coat such that the embryo is
ready to sprout when it is deposited, along with a bit of fecal matter that acts as fertilizer, far
from the parent plant. Microorganisms are often effective in breaking down hard seed coats and
are sometimes used by people as a treatment; the seeds are stored in a moist warm sandy medium
for several months under nonsterile conditions.
Stratification, also called moist-chilling, breaks down physiological dormancy, and involves the
addition of moisture to the seeds so they absorb water, and they are then subjected to a period of
moist chilling to after-ripen the embryo. Sowing in late summer and fall and allowing to
overwinter under cool conditions is an effective way to stratify seeds; some seeds respond more
favorably to periods of oscillating temperatures which are a part of the natural environment.

Leaching or the soaking in water removes chemical inhibitors in some seeds that prevent
germination. Rain and melting snow naturally accomplish this task. For seeds planted in gardens,
running water is best – if soaked in a container, 12 to 24 hours of soaking is sufficient. Soaking
longer, especially in stagnant water, can result in oxygen starvation and seed death. Seeds with
hard seed coats can be soaked in hot water to break open the impermeable cell layers that prevent
water intake.

Other methods used to assist in the germination of seeds that have dormancy include prechilling,
predrying, daily alternation of temperature, light exposure, potassium nitrate, the use of plant
growth regulators, such as gibberellins, cytokinins, ethylene, thiourea, sodium hypochlorite, and
others.[43] Some seeds germinate best after a fire. For some seeds, fire cracks hard seed coats,
while in others, chemical dormancy is broken in reaction to the presence of smoke. Liquid smoke
is often used by gardeners to assist in the germination of these species.[44]

Sterile seeds[edit]

Seeds may be sterile for few reasons: they may have been irradiated, unpollinated, cells lived
past expectancy, or bred for the purpose.

Evolution and origin of seeds[edit]

The issue of the origin of seed plants remains unsolved. However, more and more data tends to
place this origin in the middle Devonian. The description in 2004 of the proto-seed Runcaria
heinzelinii in the Givetian of Belgium is an indication of that ancient origin of seed-plants. As
with modern ferns, most land plants before this time reproduced by sending into the air spores
that would land and become whole new plants.

Taxonomists have described early "true" seeds from the upper Devonian, which probably
became the theater of their true first evolutionary radiation. With this radiation came an
evolution of seed size, shape, dispersal and eventually the radiation of gymnosperms and
angiosperms and monocotyledons and dicotyledons. Seed plants progressively became one of the
major elements of nearly all ecosystems.
Diagram of the internal structure of a dicot seed and embryo: (a) seed coat, (b) endosperm, (c)
cotyledon, (d) hypocotyl

A typical seed includes two basic parts:

1. an embryo;
2. a seed coat.

In addition, the endosperm forms a supply of nutrients for the embryo in most monocotyledons
and the endospermic dicotyledons.

Seed types[edit]

Seeds have been considered to occur in many structurally different types (Martin 1946).[7] These
are based on a number of criteria, of which the dominant one is the embryo-to-seed size ratio.
This reflects the degree to which the developing cotyledons absorb the nutrients of the
endosperm, and thus obliterate it.[7]

Six types occur amongst the monocotyledons, ten in the dicotyledons, and two in the
gymnosperms (linear and spatulate).[8] This classification is based on three characteristics:
embryo morphology, amount of endosperm and the position of the embryo relative to the
endosperm.
Diagram of a generalized dicot seed (1) versus a generalized monocot seed (2). A. Seed Coat B.
Cotyledon C. Hilum D. Plumule E. Radicle F. Endosperm

Comparison of monocotyledons and dicotyledons

Embryo[edit]

In endospermic seeds, there are two distinct regions inside the seed coat, an upper and larger
endosperm and a lower smaller embryo. The embryo is the fertilised ovule, an immature plant
from which a new plant will grow under proper conditions. The embryo has one cotyledon or
seed leaf in monocotyledons, two cotyledons in almost all dicotyledons and two or more in
gymnosperms. In the fruit of grains (caryopses) the single monocotyledon is shield shaped and
hence called a scutellum. The scutellum is pressed closely against the endosperm from which it
absorbs food, and passes it to the growing parts. Embryo descriptors include small, straight, bent,
curved and curled.

Nutrient storage[edit]

Within the seed, there usually is a store of nutrients for the seedling that will grow from the
embryo. The form of the stored nutrition varies depending on the kind of plant. In angiosperms,
the stored food begins as a tissue called the endosperm, which is derived from the mother plant
and the pollen via double fertilization. It is usually triploid, and is rich in oil or starch, and
protein. In gymnosperms, such as conifers, the food storage tissue (also called endosperm) is part
of the female gametophyte, a haploid tissue. The endosperm is surrounded by the aleurone layer
(peripheral endosperm), filled with proteinaceous aleurone grains.

Originally, by analogy with the animal ovum, the outer nucellus layer (perisperm) was referred
to as albumen, and the inner endosperm layer as vitellus. Although misleading, the term began to
be applied to all the nutrient matter. This terminology persists in referring to endospermic seeds
as "albuminous". The nature of this material is used in both describing and classifying seeds, in
addition to the embryo to endosperm size ratio. The endosperm may be considered to be
farinaceous (or mealy) in which the cells are filled with starch, as for instance cereal grains, or
not (non-farinaceous). The endosperm may also be referred to as "fleshy" or "cartilaginous" with
thicker soft cells such as coconut, but may also be oily as in Ricinus (castor oil), Croton and
Poppy. The endosperm is called "horny" when the cell walls are thicker such as date and coffee,
or "ruminated" if mottled, as in nutmeg, palms and Annonaceae.[9]

In most monocotyledons (such as grasses and palms) and some (endospermic or albuminous)
dicotyledons (such as castor beans) the embryo is embedded in the endosperm (and nucellus),
which the seedling will use upon germination. In the non-endospermic dicotyledons the
endosperm is absorbed by the embryo as the latter grows within the developing seed, and the
cotyledons of the embryo become filled with stored food. At maturity, seeds of these species
have no endosperm and are also referred to as exalbuminous seeds. The exalbuminous seeds
include the legumes (such as beans and peas), trees such as the oak and walnut, vegetables such
as squash and radish, and sunflowers. According to Bewley and Black (1978), Brazil nut storage
is in hypocotyl, this place of storage is uncommon among seeds.[10] All gymnosperm seeds are
albuminous.

Seed coat[edit]

The seed coat develops from the maternal tissue, the integuments, originally surrounding the
ovule. The seed coat in the mature seed can be a paper-thin layer (e.g. peanut) or something more
substantial (e.g. thick and hard in honey locust and coconut), or fleshy as in the sarcotesta of
pomegranate. The seed coat helps protect the embryo from mechanical injury, predators and
drying out. Depending on its development, the seed coat is either bitegmic or unitegmic.
Bitegmic seeds form a testa from the outer integument and a tegmen from the inner integument
while unitegmic seeds have only one integument. Usually parts of the testa or tegmen form a
hard protective mechanical layer. The mechanical layer may prevent water penetration and
germination. Amongst the barriers may be the presence of lignified sclereids.[11]

The outer integument has a number of layers, generally between four and eight organised into
three layers: (a) outer epidermis, (b) outer pigmented zone of two to five layers containing tannin
and starch, and (c) inner epidermis. The endotegmen is derived from the inner epidermis of the
inner integument, the exotegmen from the outer surface of the inner integument. The endotesta
is derived from the inner epidermis of the outer integument, and the outer layer of the testa from
the outer surface of the outer integument is referred to as the exotesta. If the exotesta is also the
mechanical layer, this is called an exotestal seed, but if the mechanical layer is the endotegmen,
then the seed is endotestal. The exotesta may consist of one or more rows of cells that are
elongated and pallisade like (e.g. Fabaceae), hence 'palisade exotesta'.[12][13]

In addition to the three basic seed parts, some seeds have an appendage, an aril, a fleshy
outgrowth of the funicle (funiculus), (as in yew and nutmeg) or an oily appendage, an elaiosome
(as in Corydalis), or hairs (trichomes). In the latter example these hairs are the source of the
textile crop cotton. Other seed appendages include the raphe (a ridge), wings, caruncles (a soft
spongy outgrowth from the outer integument in the vicinity of the micropyle), spines, or
tubercles.

A scar also may remain on the seed coat, called the hilum, where the seed was attached to the
ovary wall by the funicle. Just below it is a small pore, representing the micropyle of the ovule.

Size and seed set[edit]

A collection of various vegetable and herb seeds

Seeds are very diverse in size. The dust-like orchid seeds are the smallest, with about one million
seeds per gram; they are often embryonic seeds with immature embryos and no significant
energy reserves. Orchids and a few other groups of plants are mycoheterotrophs which depend
on mycorrhizal fungi for nutrition during germination and the early growth of the seedling. Some
terrestrial orchid seedlings, in fact, spend the first few years of their lives deriving energy from
the fungi and do not produce green leaves.[14] At over 20 kg, the largest seed is the coco de mer.
Plants that produce smaller seeds can generate many more seeds per flower, while plants with
larger seeds invest more resources into those seeds and normally produce fewer seeds. Small
seeds are quicker to ripen and can be dispersed sooner, so fall blooming plants often have small
seeds. Many annual plants produce great quantities of smaller seeds; this helps to ensure at least
a few will end in a favorable place for growth. Herbaceous perennials and woody plants often
have larger seeds; they can produce seeds over many years, and larger seeds have more energy
reserves for germination and seedling growth and produce larger, more established seedlings
after germination.[15][16]

Functions[edit]
Seeds serve several functions for the plants that produce them. Key among these functions are
nourishment of the embryo, dispersal to a new location, and dormancy during unfavorable
conditions. Seeds fundamentally are means of reproduction, and most seeds are the product of
sexual reproduction which produces a remixing of genetic material and phenotype variability on
which natural selection acts.

Embryo nourishment[edit]

Seeds protect and nourish the embryo or young plant. They usually give a seedling a faster start
than a sporeling from a spore, because of the larger food reserves in the seed and the
multicellularity of the enclosed embryo.

Dispersal[edit]

Main article: Seed dispersal

Unlike animals, plants are limited in their ability to seek out favorable conditions for life and
growth. As a result, plants have evolved many ways to disperse their offspring by dispersing
their seeds (see also vegetative reproduction). A seed must somehow "arrive" at a location and be
there at a time favorable for germination and growth. When the fruits open and release their
seeds in a regular way, it is called dehiscent, which is often distinctive for related groups of
plants; these fruits include capsules, follicles, legumes, silicles and siliques. When fruits do not
open and release their seeds in a regular fashion, they are called indehiscent, which include the
fruits achenes, caryopsis, nuts, samaras, and utricles.[17]

By wind (anemochory)[edit]

Dandelion seeds are contained within achenes, which can be carried long distances by the wind.

The seed pod of milkweed (Asclepias syriaca)

 Some seeds (e.g., pine) have a wing that aids in wind dispersal.
 The dustlike seeds of orchids are carried efficiently by the wind.
 Some seeds (e.g. milkweed, poplar) have hairs that aid in wind dispersal.[18]

Other seeds are enclosed in fruit structures that aid wind dispersal in similar ways:
  Dandelion achenes have hairs.
 Maple samaras have two wings.

By water (hydrochory)[edit]

 Some plants, such as Mucuna and Dioclea, produce buoyant seeds termed sea-beans or drift
seeds because they float in rivers to the oceans and wash up on beaches.[19]

By animals (zoochory)[edit]

 Seeds (burrs) with barbs or hooks (e.g. acaena, burdock, dock) which attach to animal fur or
feathers, and then drop off later.
 Seeds with a fleshy covering (e.g. apple, cherry, juniper) are eaten by animals (birds, mammals,
reptiles, fish) which then disperse these seeds in their droppings.
 Seeds (nuts) are attractive long-term storable food resources for animals (e.g. acorns, hazelnut,
walnut); the seeds are stored some distance from the parent plant, and some escape being
eaten if the animal forgets them.

Myrmecochory is the dispersal of seeds by ants. Foraging ants disperse seeds which have
appendages called elaiosomes[20] (e.g. bloodroot, trilliums, acacias, and many species of
Proteaceae). Elaiosomes are soft, fleshy structures that contain nutrients for animals that eat
them. The ants carry such seeds back to their nest, where the elaiosomes are eaten. The
remainder of the seed, which is hard and inedible to the ants, then germinates either within the
nest or at a removal site where the seed has been discarded by the ants.[21] This dispersal
relationship is an example of mutualism, since the plants depend upon the ants to disperse seeds,
while the ants depend upon the plants seeds for food. As a result, a drop in numbers of one
partner can reduce success of the other. In South Africa, the Argentine ant (Linepithema humile)
has invaded and displaced native species of ants. Unlike the native ant species, Argentine ants do
not collect the seeds of Mimetes cucullatus or eat the elaiosomes. In areas where these ants have
invaded, the numbers of Mimetes seedlings have dropped.[22]

Dormancy[edit]

Main article: 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 (e.g. late frosts, drought, herbivory) does not result in the
death of all offspring of a plant (bet-hedging).[23] 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.[24] 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.[25]

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.[26]

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.[27]
 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.[28] 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.[29]

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.[30][31]
 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.[30] 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.[30]
 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.
o 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.[32]
 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.[31]
 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.[33]

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.[34]

Persistence and seed banks[edit]

Further information: Seed dormancy

Germination[edit]
Main articles: Seedling and Germination
Germinating sunflower seedlings

Seed germination is a process by which a seed embryo develops into a seedling. It involves the
reactivation of the metabolic pathways that lead to growth and the emergence of the radicle or
seed root and plumule or shoot. The emergence of the seedling above the soil surface is the next
phase of the plant's growth and is called seedling establishment.[35]

Three fundamental conditions must exist before germination can occur. (1) The embryo must be
alive, called seed viability. (2) Any dormancy requirements that prevent germination must be
overcome. (3) The proper environmental conditions must exist for germination.

Seed viability is the ability of the embryo to germinate and is affected by a number of different
conditions. Some plants do not produce seeds that have functional complete embryos, or the seed
may have no embryo at all, often called empty seeds. Predators and pathogens can damage or kill
the seed while it is still in the fruit or after it is dispersed. Environmental conditions like flooding
or heat can kill the seed before or during germination. The age of the seed affects its health and
germination ability: since the seed has a living embryo, over time cells die and cannot be
replaced. Some seeds can live for a long time before germination, while others can only survive
for a short period after dispersal before they die.

Seed vigor is a measure of the quality of seed, and involves the viability of the seed, the
germination percentage, germination rate and the strength of the seedlings produced.[36]

The germination percentage is simply the proportion of seeds that germinate from all seeds
subject to the right conditions for growth. The germination rate is the length of time it takes for
the seeds to germinate. Germination percentages and rates are affected by seed viability,
dormancy and environmental effects that impact on the seed and seedling. In agriculture and
horticulture quality seeds have high viability, measured by germination percentage plus the rate
of germination. This is given as a percent of germination over a certain amount of time, 90%
germination in 20 days, for example. 'Dormancy' is covered above; many plants produce seeds
with varying degrees of dormancy, and different seeds from the same fruit can have different
degrees of dormancy.[37] It's possible to have seeds with no dormancy if they are dispersed right
away and do not dry (if the seeds dry they go into physiological dormancy). There is great
variation amongst plants and a dormant seed is still a viable seed even though the germination
rate might be very low.
Environmental conditions affecting seed germination include; water, oxygen, temperature and
light.

Three distinct phases of seed germination occur: water imbibition; lag phase; and radicle
emergence.

In order for the seed coat to split, the embryo must imbibe (soak up water), which causes it to
swell, splitting the seed coat. However, the nature of the seed coat determines how rapidly water
can penetrate and subsequently initiate germination. The rate of imbibition is dependent on the
permeability of the seed coat, amount of water in the environment and the area of contact the
seed has to the source of water. For some seeds, imbibing too much water too quickly can kill the
seed. For some seeds, once water is imbibed the germination process cannot be stopped, and
drying then becomes fatal. Other seeds can imbibe and lose water a few times without causing ill
effects, but drying can cause secondary dormancy.

Repair of DNA damage[edit]

During seed dormancy, often associated with unpredictable and stressful environments, DNA
damage accumulates as the seeds age.[38][39][40] In rye seeds, the reduction of DNA integrity due to
damage is associated with loss of seed viability during storage.[38] Upon germination, seeds of
Vicia faba undergo DNA repair.[39] A plant DNA ligase that is involved in repair of single- and
double-strand breaks during seed germination is an important determinant of seed longevity.[41]
Also, in Arabidopsis seeds, the activities of the DNA repair enzymes Poly ADP ribose
polymerases (PARP) are likely needed for successful germination.[42] Thus DNA damages that
accumulate during dormancy appear to be a problem for seed survival, and the enzymatic repair
of DNA damages during germination appears to be important for seed viability.

Inducing germination[edit]

A number of different strategies are used by gardeners and horticulturists to break seed
dormancy.

Scarification allows water and gases to penetrate into the seed; it includes methods to physically
break the hard seed coats or soften them by chemicals, such as soaking in hot water or poking
holes in the seed with a pin or rubbing them on sandpaper or cracking with a press or hammer.
Sometimes fruits are harvested while the seeds are still immature and the seed coat is not fully
developed and sown right away before the seed coat become impermeable. Under natural
conditions, seed coats are worn down by rodents chewing on the seed, the seeds rubbing against
rocks (seeds are moved by the wind or water currents), by undergoing freezing and thawing of
surface water, or passing through an animal's digestive tract. In the latter case, the seed coat
protects the seed from digestion, while often weakening the seed coat such that the embryo is
ready to sprout when it is deposited, along with a bit of fecal matter that acts as fertilizer, far
from the parent plant. Microorganisms are often effective in breaking down hard seed coats and
are sometimes used by people as a treatment; the seeds are stored in a moist warm sandy medium
for several months under nonsterile conditions.
Stratification, also called moist-chilling, breaks down physiological dormancy, and involves the
addition of moisture to the seeds so they absorb water, and they are then subjected to a period of
moist chilling to after-ripen the embryo. Sowing in late summer and fall and allowing to
overwinter under cool conditions is an effective way to stratify seeds; some seeds respond more
favorably to periods of oscillating temperatures which are a part of the natural environment.

Leaching or the soaking in water removes chemical inhibitors in some seeds that prevent
germination. Rain and melting snow naturally accomplish this task. For seeds planted in gardens,
running water is best – if soaked in a container, 12 to 24 hours of soaking is sufficient. Soaking
longer, especially in stagnant water, can result in oxygen starvation and seed death. Seeds with
hard seed coats can be soaked in hot water to break open the impermeable cell layers that prevent
water intake.

Other methods used to assist in the germination of seeds that have dormancy include prechilling,
predrying, daily alternation of temperature, light exposure, potassium nitrate, the use of plant
growth regulators, such as gibberellins, cytokinins, ethylene, thiourea, sodium hypochlorite, and
others.[43] Some seeds germinate best after a fire. For some seeds, fire cracks hard seed coats,
while in others, chemical dormancy is broken in reaction to the presence of smoke. Liquid smoke
is often used by gardeners to assist in the germination of these species.[44]

Sterile seeds[edit]

Seeds may be sterile for few reasons: they may have been irradiated, unpollinated, cells lived
past expectancy, or bred for the purpose.

Evolution and origin of seeds[edit]

The issue of the origin of seed plants remains unsolved. However, more and more data tends to
place this origin in the middle Devonian. The description in 2004 of the proto-seed Runcaria
heinzelinii in the Givetian of Belgium is an indication of that ancient origin of seed-plants. As
with modern ferns, most land plants before this time reproduced by sending into the air spores
that would land and become whole new plants.

Taxonomists have described early "true" seeds from the upper Devonian, which probably
became the theater of their true first evolutionary radiation. With this radiation came an
evolution of seed size, shape, dispersal and eventually the radiation of gymnosperms and
angiosperms and monocotyledons and dicotyledons. Seed plants progressively became one of the
major elements of nearly all ecosystems.

Economic importance[edit]
Phaseolus vulgaris (common bean or green bean) seeds are diverse in size, shape, and color.

Seed market[edit]

In the United States farmers spent $22 billion on seeds in 2018, a 35 percent increase since 2010.
DowDuPont and Monsanto account for 72 percent of corn and soybean seed sales in the U.S.
with the average price of a bag of GMO corn seed is priced at $270.[45]

Edible seeds[edit]

Further information: List of edible seeds

Many seeds are edible and the majority of human calories comes from seeds,[46] especially from
cereals, legumes and nuts. Seeds also provide most cooking oils, many beverages and spices and
some important food additives. In different seeds the seed embryo or the endosperm dominates
and provides most of the nutrients. The storage proteins of the embryo and endosperm differ in
their amino acid content and physical properties. For example, the gluten of wheat, important in
providing the elastic property to bread dough is strictly an endosperm protein.

Seeds are used to propagate many crops such as cereals, legumes, forest trees, turfgrasses, and
pasture grasses. Particularly in developing countries, a major constraint faced is the inadequacy
of the marketing channels to get the seed to poor farmers.[47] Thus the use of farmer-retained seed
remains quite common.

Seeds are also eaten by animals (seed predation), and are also fed to livestock or provided as
birdseed.

Poison and food safety[edit]

While some seeds are edible, others are harmful, poisonous or deadly.[48] Plants and seeds often
contain chemical compounds to discourage herbivores and seed predators. In some cases, these
compounds simply taste bad (such as in mustard), but other compounds are toxic or break down
into toxic compounds within the digestive system. Children, being smaller than adults, are more
susceptible to poisoning by plants and seeds.[49]
A deadly poison, ricin, comes from seeds of the castor bean. Reported lethal doses are anywhere
from two to eight seeds,[50][51] though only a few deaths have been reported when castor beans
have been ingested by animals.[52]

In addition, seeds containing amygdalin – apple, apricot, bitter almond,[53] peach, plum, cherry,
quince, and others – when consumed in sufficient amounts, may cause cyanide poisoning.[53][54]
Other seeds that contain poisons include annona, cotton, custard apple, datura, uncooked durian,
golden chain, horse-chestnut, larkspur, locoweed, lychee, nectarine, rambutan, rosary pea, sour
sop, sugar apple, wisteria, and yew.[50][55] The seeds of the strychnine tree are also poisonous,
containing the poison strychnine.

The seeds of many legumes, including the common bean (Phaseolus vulgaris), contain proteins
called lectins which can cause gastric distress if the beans are eaten without cooking. The
common bean and many others, including the soybean, also contain trypsin inhibitors which
interfere with the action of the digestive enzyme trypsin. Normal cooking processes degrade
lectins and trypsin inhibitors to harmless forms.[56]

Please see the category plant toxins for further relevant articles.

Other uses[edit]

Cotton fiber grows attached to cotton plant seeds. Other seed fibers are from kapok and
milkweed.

Many important nonfood oils are extracted from seeds. Linseed oil is used in paints. Oil from
jojoba and crambe are similar to whale oil.

Seeds are the source of some medicines including castor oil, tea tree oil and the quack cancer
drug Laetrile.

Many seeds have been used as beads in necklaces and rosaries including Job's tears, Chinaberry,
rosary pea, and castor bean. However, the latter three are also poisonous.

Other seed uses include:

 Seeds once used as weights for balances.

 Seeds used as toys by children, such as for the game Conkers.
 Resin from Clusia rosea seeds used to caulk boats.
 Nematicide from milkweed seeds.
 Cottonseed meal used as animal feed and fertilizer.

Seed records[edit]
The massive fruit of the coco de mer

 The oldest viable carbon-14-dated seed that has grown into a plant was a Judean date palm
seed about 2,000 years old, recovered from excavations at Herod the Great's palace on Masada
in Israel. It was germinated in 2005.[57] (A reported regeneration of Silene stenophylla (narrow-
leafed campion) from material preserved for 31,800 years in the Siberian permafrost was
achieved using fruit tissue, not seed.[58][59])
 The largest seed is produced by the coco de mer, or "double coconut palm", Lodoicea maldivica.
The entire fruit may weigh up to 23 kilograms (50 pounds) and usually contains a single seed.[60]
 The smallest seeds are produced by epiphytic orchids. They are only 85 micrometers long, and
weigh 0.81 micrograms. They have no endosperm and contain underdeveloped embryos.[61]
 The earliest fossil seeds are around 365 million years old from the Late Devonian of West
Virginia. The seeds are preserved immature ovules of the plant Elkinsia polymorpha.[62]

In religion[edit]
The Book of Genesis in the Old Testament begins with an explanation of how all plant forms
began:

And God said, Let the earth bring forth grass, the herb yielding seed, and the fruit tree yielding
fruit after his kind, whose seed is in itself, upon the earth: and it was so. And the earth brought
forth grass, and herb yielding seed after its kind, and the tree yielding fruit, whose seed was in
itself, after its kind: and God saw that it was good. And the evening and the morning were the
third day.[63]

The Quran speaks of seed germination thus:

It is Allah Who causeth the seed-grain and the date-stone to split and sprout. He causeth the
living to issue from the dead, and He is the one to cause the dead to issue from the living. That is
Allah: then how are ye deluded away from the truth?[
References[edit]
1. ^ Cain M.D., Shelton M.G. (2001). "Twenty years of natural loblolly and shortleaf pine seed
production on the Crossett Experimental Forest in southeastern Arkansas". Southern Journal of
Applied Forestry. 25 (1): 40–45.
2. ^ Galili G; Kigel J (1995). "Chapter One". Seed development and germination. New York: M.
Dekker. ISBN 978-0-8247-9229-9.
3. ^ Raven, Peter H., Ray Franklin Evert, and Helena Curtis. 1981. Biology of plants. New York:
Worth Publishers. p. 410.
4. ^ Rost, Thomas L.; Weier, T. Elliot; Weier, Thomas Elliot (1979). Botany: a brief introduction to
plant biology. New York: Wiley. p. 319. ISBN 978-0-471-02114-8.
5. ^ Filonova LH; Bozhkov PV; von Arnold S (February 2000). "Developmental pathway of somatic
embryogenesis in Picea abies as revealed by time-lapse tracking". J Exp Bot. 51 (343): 249–264.
doi:10.1093/jexbot/51.343.249. PMID 10938831. Archived from the original on 2008-12-07.
6. ^ "Seed shape". anbg.gov.au. Archived from the original on 2014-02-26.
7. ^ Jump up to: a b The Seed Biology Place, Gerhard Leubner Lab, Royal Holloway, University of
London, archived from the original on 24 September 2015, retrieved 13 October 2015
8. ^ Baskin, Carol C.; Baskin, Jerry M. (2001). Carol C. Baskin, Jerry M. Baskin. Seeds: Ecology,
Biogeography, and Evolution of Dormancy and Germination. Elsevier, 2001. google.ca. p. 27.
ISBN 978-0-12-080263-0.
9. ^ "The Encyclopædia Britannica, 9th ed. (1888) vol. 4". google.ca. 1888. p. 155.
10. ^ Bewley & Black (1978) Physiology and Biochemistry of Seeds in Relation to Germination,
pag.11
11. ^ "Sinauer Associates, Inc., Publishers". 5e.plantphys.net. Archived from the original on 22
January 2014. Retrieved 7 May 2018.
12. ^ "plant_anatomy Term "seed coat epidermis" (PO:0006048)". gramene.org. Archived from the
original on 2014-02-03.
13. ^ Rudall, Paula J. (2007). 6 – Seed and fruit – University Publishing Online – Paula J. Rudall.
Anatomy of Flowering Plants: An Introduction to Structure and Development. Third edition.
Cambridge University Press. doi:10.1017/CBO9780511801709. ISBN 978-0-521-69245-8.
14. ^ Smith, Welby R. 1993. Orchids of Minnesota. Minneapolis: University of Minnesota Press. p.
8.
15. ^ Kosinki, Igor (2007). "Long-term variability in seed size and seedling establishment of
Maianthemum bifolium". Plant Ecology. 194 (2): 149–156. doi:10.1007/s11258-007-9281-1.
16. ^ Shannon DA; Isaac L; Brockman FE (February 1996). "Assessment of hedgerow species for
seed size, stand establishment and seedling height". Agroforestry Systems. 35 (1): 95–110.
doi:10.1007/BF02345331.
17. ^ Jones, Samuel B., and Arlene E. Luchsinger. 1979. Plant systematics. McGraw-Hill series in
organismic biology. New York: McGraw-Hill. p. 195.
18. ^ Morhardt, Sia; Morhardt, Emil; Emil Morhardt, J. (2004). California desert flowers: an
introduction to families, genera, and species. Berkeley: University of California Press. p. 24.
ISBN 978-0-520-24003-2.
19. ^ "www.seabean.com – Sea-Beans and Drift Seeds". seabean.com. Archived from the original on
2006-07-11.
20. ^ Marinelli J (1999). "Ants – The astonishing intimacy between ants & plants". Plants &
Gardens News. 14 (1). Archived from the original on 2006-08-18.
21. ^ Ricklefs, Robert E. (1993) The Economy of Nature, 3rd ed., p. 396. (New York: W.H.
Freeman). ISBN 0-7167-2409-X.
 22. ^ Bond, W.J.; P. Slingsby (1984). "Collapse of an ant-plant mutualism: The Argentine ant,
Iridomyrmex humilis and myrmecochorous Proteaceae". Ecology. 65 (4): 1031–1037.
doi:10.2307/1938311. JSTOR 1938311.
23. ^ Eira MTS, Caldas LS (2000) Seed dormancy and germination as concurrent processes. Rev
Bras Fisiol Vegetal 12:85–104

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 with in the motherplant.
The formation of the seed completes the process of reproduction in plants (started with the development of flowers
and pollination), with the embryo developed from the zygote and the seed coat from the integuments of the ovule.
This process starts with double fertilization in angiosperms and it involves the fusion of the egg and sperm nuclei into
a zygote.
The second part of this process is the fusion of the polar nuclei with a second sperm cell nucleus, thus forming a
primary endosperm.
Right after fertilization the zygote is mostly inactive but the primary endosperm divides rapidly to form the endosperm
tissue.
This tissue becomes the food that the young plant will consume until the roots have developed after germination, or it
develops into a hard seed coat.
The seed, which is an embryo with two points of growth (one of which forms the stems the other the roots) is
enclosed in a seed coat with some food reserves.
In gymnosperms the two sperm cells transferred from the pollen do not develop seed by double fertilization but
instead only one sperm fertilizes the egg while the other is not used.
The seed is composed of the embryo (the result of fertilization) and tissue from the mother plant, which also form a
cone around the seed in coniferous plants like Pine and Spruce.
The new seed is formed in plant structures called fruits.
Plants have evolved many ways to disperse and spread the population through their seeds.

Seed
PLANT REPRODUCTIVE PART

WRITTEN BY:

 Hans Lambers
See Article History
 seed storage in vascular seed plantsVideo presentation describing the differences in seed storage between angiosperms and
gymnosperms.Encyclopædia Britannica, Inc.

Seed, the characteristic reproductive body of both angiosperms (flowering plants)

and gymnosperms (e.g., conifers, cycads, and ginkgos). Essentially, a seed consists of a
miniature undeveloped plant (the embryo), which, alone or in the company of stored food for
its early development after germination, is surrounded by a protective coat (the testa).
Frequently small in size and making negligible demands upon their environment, seeds are
eminently suited to perform a wide variety of functions the relationships of which are not
always obvious: multiplication, perennation (surviving seasons of stress such as
winter), dormancy (a state of arrested development), and dispersal. Pollination and the
“seed habit” are considered the most important factors responsible for the overwhelming
evolutionary success of the flowering plants, which number more than 300,000 species.
magnolia fruit and seedFruit and seeds of the southern magnolia (Magnolia grandiflora). The fruit is an aggregate of follicles, and
each follicle bears a single red seed.© Maljalen/Dreamstime.com

The superiority of dispersal by means of seeds over the more primitive method involving
single-celled spores, lies mainly in two factors: the stored reserve of nutrient material that
gives the new generation an excellent growing start and the seed’s multicellular structure.
The latter factor provides ample opportunity for the development of adaptations for
dispersal, such as plumes for wind dispersal, barbs, and others.
 seed dispersalSeeds and their dispersal mechanisms.Encyclopædia Britannica, Inc.

Economically, seeds are important primarily because they are sources of a variety of
foods—for example, the cereal grains, such as wheat, rice, and corn (maize); the seeds
of beans, peas, peanuts, soybeans, almonds, sunflowers, hazelnuts, walnuts, pecans,
and Brazil nuts. Other useful products provided by seeds are
abundant. Oils for cooking, margarine production, painting, and lubrication are available
from the seeds of flax, rape, cotton, soybean, poppy, castor
bean, coconut, sesame, safflower, sunflower, and various cereal grains. Essential oils are
obtained from such sources as juniper “berries,” used in gin manufacture. Stimulants are
obtained from such sources as the seeds of coffee, kola, guarana, and cocoa. Spices—
from mustard and nutmeg seeds; from the aril (“mace”) covering the nutmeg seed; from the
seeds and fruits of anise, cumin, caraway, dill, vanilla, black pepper, allspice, and others—
form a large group of economic products.
 


caraway seedsCaraway seeds.DO'Neil

castor bean seedsCastor bean seeds used to make oil cakes.Brian Prechtel/U.S. Department of Agriculture (Image number: K9200-2)
 Advertisement

The Nature Of Seeds

Angiosperm seeds
In the typical flowering plant, or angiosperm, seeds are formed from bodies
called ovules contained in the ovary, or basal part of the female plant structure, the pistil.
The mature ovule contains in its central part a region called the nucellus that in turn
contains an embryo sac with eight nuclei, each with one set of chromosomes (i.e., they are
haploid nuclei). The two nuclei near the centre are referred to as polar nuclei; the egg cell,
or oosphere, is situated near the micropylar (“open”) end of the ovule.

Diagram of a typical flowering plant (angiosperm).Encyclopædia Britannica, Inc.

Facts Matter. Support the truth and unlock all of Britannica’s content.Start Your Free Trial
Today

With very few exceptions (e.g., the dandelion), development of the ovule into a seed is
dependent upon fertilization, which in turn follows pollination. Pollen grains that land on the
receptive upper surface (stigma) of the pistil will germinate, if they are of the same species,
and produce pollen tubes, each of which grows down within the style (the upper part of the
pistil) toward an ovule. The pollen tube has three haploid nuclei, one of them, the so-
called vegetative, or tube, nucleus seems to direct the operations of the growing structure.
The other two, the generative nuclei, can be thought of as nonmotile sperm cells. After
reaching an ovule and breaking out of the pollen tube tip, one generative nucleus unites
with the egg cell to form a diploid zygote (i.e., a fertilized egg with two complete sets
of chromosomes, one from each parent). The zygote undergoes a limited number
of divisions and gives rise to an embryo. The other generative nucleus fuses with the two
polar nuclei to produce a triploid (three sets of chromosomes) nucleus, which divides
repeatedly before cell-wall formation occurs. This process gives rise to the
triploid endosperm, a nutrient tissue that contains a variety of storage materials—such
as starch, sugars, fats, proteins, hemicelluloses, and phytate (a phosphate reserve).
The events just described constitute what is called the double-fertilization process, one of
the characteristic features of all flowering plants. In the orchids and in some other plants
with minute seeds that contain no reserve materials, endosperm formation is completely
suppressed. In other cases it is greatly reduced, but the reserve materials are present
elsewhere—e.g., in the cotyledons, or seed leaves, of the embryo, as in beans, lettuce,
and peanuts, or in a tissue derived from the nucellus, the perisperm, as in coffee. Other
seeds, such as those of beets, contain both perisperm and endosperm. The seed coat, or
testa, is derived from the one or two protective integuments of the ovule. The ovary, in the
simplest case, develops into a fruit. In many plants, such as grasses and lettuce, the outer
integument and ovary wall are completely fused, so seed and fruit form one entity; such
seeds and fruits can logically be described together as “dispersal units,” or diaspores. More
often, however, the seeds are discrete units attached to the placenta on the inside of the
fruit wall through a stalk, or funiculus.
The hilum of a liberated seed is a small scar marking its former place of attachment. The
short ridge (raphe) that sometimes leads away from the hilum is formed by the fusion of
seed stalk and testa. In many seeds, the micropyle of the ovule also persists as a small
opening in the seed coat. The embryo, variously located in the seed, may be very small (as
in buttercups) or may fill the seed almost completely (as in roses and plants of the mustard
family). It consists of a root part, or radicle, a prospective shoot (plumule or epicotyl), one or
more cotyledons (one or two in flowering plants, several in Pinus and other gymnosperms),
and a hypocotyl, which is a region that connects radicle and plumule. A classification of
seeds can be based on size and position of the embryo and on the proportion of embryo to
storage tissue; the possession of either one or two cotyledons is considered crucial in
recognizing two main groups of flowering plants, the monocotyledons and
the eudicotyledons.

epigeal germinationTime-lapse video of the epigeal (cotyledons emerge aboveground) germination of a dwarf French bean
(Phaseolus vulgaris ‘Borlotto Firetongue'), filmed over two weeks.Video by Neil Bromhall; music, Telemann Trio/Musopen.org (A
Britannica Publishing Partner)

hypogeal germinationTime-lapse video of the hypogeal (cotyledons remain belowground) germination of runner beans (Phaseolus
coccineus ‘Enorma'), filmed over a three-week period.Video by Neil Bromhall; music, Paul Pitman/Musopen.org (A Britannica
Publishing Partner)

Seedlings, arising from embryos in the process of germination, are classified as epigeal
(cotyledons aboveground, usually green and capable of photosynthesis) and hypogeal
(cotyledons belowground). Particularly in the monocots, special absorbing organs may
develop that mobilize the reserve materials and withdraw them from the endosperm; e.g., in
grasses, the cotyledon has been modified into an enzyme-secreting scutellum (“shield”)
between embryo and endosperm.

Seed

QUICK FACTS

KEY PEOPLE

 W. Atlee Burpee

RELATED TOPICS
  Soil seed bank
 Peanut
 Cottonseed
 Soybean
 Flaxseed
 Mast seeding
 Aril
 Endosperm
 Caruncle
 Seed bank
Gymnosperm seeds
In gymnosperms (plants with “naked seeds”—such as conifers, cycads, and ginkgo), the
ovules are not enclosed in an ovary but lie exposed on leaflike structures,
the megasporophylls. A long time span usually separates pollination and fertilization, and
the ovules begin to develop into seeds long before fertilization has been accomplished; in
some cases, in fact, fertilization does not occur until the ovules (“seeds”) have been shed
from the tree. In the European, or Scots, pine (Pinus sylvestris), for example, the female
cones (essentially collections of megasporophylls) begin to develop in winter and are ready
to receive pollen from the male cones in spring. During the first growing season, the pollen
tube grows slowly through the nucellus, while within the ovule the megaspore nucleus,
through a series of divisions, gives rise to a collection of some 2,000 nuclei, which are then
individually enclosed by walls to form a structure called the
female gametophyte or prothallus. At the micropylar end of the ovule,
several archegonia (bottle-shaped female organs) develop, each containing an oosphere
(“egg”). The pollen tube ultimately penetrates the neck of one of the archegonia. Not until
the second growing season, however, does the nucleus of one of the male cells in the tube
unite with the oosphere nucleus. Although more than one archegonium may be fertilized,
only one gives rise to a viable embryo. During the latter’s development, part of the
prothallus is broken down and used. The remainder, referred to as endosperm, surrounds
the embryo; it is mobilized later, during germination of the seed, a process that occurs
without delay when the seeds are liberated from the female cone during the third year after
their initiation.
 pinyon pinePinecone and exposed seeds of the pinyon pine (Pinus edulis). Pinyon pines are gymnosperms and bear their edible
seeds, known as pine nuts, in protective cones instead of fruit.Doak Heyser—iStock/Thinkstock

Form And Function

Seed size

In the Late Carboniferous Period (about 315.2 million to 298.9 million years ago),
some seed ferns produced large seeds (12 × 6 cm [5 × 2 inches] in Pachytesta incrassata).
This primitive ancestral condition of large seeds is reflected in certain gymnosperms (Cycas
circinalis, 5.5 × 4 cm [2.2 × 1.6 inches]; Araucaria bidwillii, 4.5 × 3.5 cm [1.8 × 1.4 inches])
and also in some tropical rainforest trees with nondormant water-rich seeds (Mora excelsa,
12 × 7 cm [4.7 × 2.8 inches]). The “double coconut” palm Lodoicea maldivica represents the
extreme, with seeds weighing up to 27 kg (about 60 pounds). Herbaceous nontropical
flowering plants usually have seeds weighing in the range of about 0.0001 to 0.01 gram.
Within a given family (e.g., the pea family, Fabaceae), seed size may vary greatly; in others
it is consistently large or small, justifying the recognition of “megaspermous” families
(e.g., beech, nutmeg, palm, and soursop families) and “microspermous” ones (e.g.,
milkweed, daisy, heather, nettle, and willow families).
 The smallest known seeds, devoid of food reserves, are found in orchids, mycoheterotrophs
(nongreen plants that absorb nutrients from dead organic matter and live symbiotically
with mycorrizal fungi—e.g., Indian pipe, Monotropa; coral root, Corallorhiza), carnivorous
plants (sundews, pitcher plants), and total parasites (members of the
families Rafflesiaceae and Orobanchaceae, or broomrapes, which have seeds weighing as
little as 0.001 mg—about 3.5 hundred-millionths of an ounce). Clearly, seed size is related
to lifestyle. Total parasites obtain food from their host, even in their early growth stages, and
young orchids are mycoheterotrophs that receive assistance in absorbing nutrients from
mycorrhizal fungi that are associated closely with their roots. In both cases only very small
seeds that lack endosperm are produced. Dodders (Cuscuta)
and mistletoes (Viscum, Phoradendron, Amyema) live independently when very young and
accordingly have relatively large seeds.
Many plant species possess seeds of remarkably uniform size, useful as beads (e.g., Abrus
precatorius) or units of weight—one carat of weight once corresponded with one seed of
the carob tree, Ceratonia siliqua. In wheat and many other plants, average seed size does
not depend on planting density, showing that seed size is under rather strict genetic control.
This does not necessarily preclude significant variation among individual seeds; in peas, for
example, the seeds occupying the central region of the pod are the largest, probably as the
result of competition for nutrients between developing ovules on the placenta. Striking
evolutionary changes in seed size, inadvertently created by humans, have occurred in
the weed known as gold-of-pleasure (Camelina sativa), which grows in flax fields. The
customary winnowing of flax seeds selects forms of C. sativa whose seeds are blown over
the same distance as flax seeds in the operation, thus staying with their “models.”
Consequently, C. sativa seeds in the south of Russia now mimic the relatively thick, heavy
seeds of the oil flax that is grown there, whereas in the northwest they resemble the flat,
thin seeds of the predominant fibre flax.

SIMILAR TOPICS

 Fruit
 Plant development
 Alternation of generations
 Germination
 Perennial
 Biennial
 Annual
Seed size and predation

Seeds form the main source of food for many birds, rodents, ants, and beetles. Harvester
ants of the genus Veromessor, for example, exact a toll of about 15,000,000 seeds per acre
(37,050,000 seeds per hectare) per year from the Sonoran Desert of the southwestern
United States. In view of the enormous size range of the predators, which include
minute weevil and bruchid-beetle larvae that attack the seeds internally, evolutionary
“manipulation” of seed size by a plant species cannot in itself be effective in completely
avoiding seed attack. With predation inescapable, however, it must be advantageous for a
plant species to invest the total reproductive effort in a large number of very small units
(seeds) rather than in a few big ones. The mean seed weight of those 13 species of Central
American woody legumes vulnerable to bruchid attack is 0.26 gram (0.009 ounce). In
contrast, the mean seed weight of the 23 species invulnerable by virtue of toxic
seed constituents is 3 grams (0.1 ounce).

READ MORE ON THIS TOPIC

plant reproductive system: Seed plants

In the two great groups of seed plants, gymnosperms and angiosperms, the sporophyte is the
dominant phase in the life cycle, as it is also…

Seed size and germination

Ecologically, seed size is also important in the breaking of dormancy. Being small, a seed
can only “sample” that part of the environment immediately adjacent to it, which is not
necessarily representative of the generally prevailing conditions. For successful seedling
establishment, there is clearly a risk in “venturing out” in adverse conditions. The
development in seeds of mechanisms acting as “integrating rain gauges” should be
considered in that light (see below).
The shape of dispersal units
Apart from the importance of shape as a factor in determining the mode of dispersal (e.g.,
wind dispersal of winged seeds, animal dispersal of spiny fruits), shape also counts when
the seed or diaspore is seen as a landing device. The flatness of the enormous
tropical Mora seeds prevents rolling and effectively restricts germination to the spot where
they land. In contrast, Eusideroxylon zwageri does not grow on steep slopes, because its
heavy fruits roll downhill. The grains of the grass Panicum turgidum, which have a flat and a
round side, germinate much better when the flat rather than the convex side lies in contact
with wet soil. In very small seeds, the importance of shape can be judged only by taking into
account soil clod size and microtopography of the soils onto which they are dropped. The
rounded seeds of cabbage species, for example, tend to roll into crevices, whereas the
reticulate ones of lamb’s quarters (Chenopodium album) often stay in the positions in which
they first fall. Several seeds have appendages (awns, bristles) that promote germination by
aiding in orientation and self-burial. In one study, for example, during a six-month period,
awned grains of Danthonia penicillata gave rise to 12 times as many established seedlings
as de-awned ones.
Polymorphism of seeds and fruits

Some plant species produce two or more sharply defined types of seeds that differ in
appearance, colour, shape, size, internal structure, or dormancy. In common spurry
(Spergula arvensis), for example, the seed coat (part of the mother plant) may be either
smooth or papillate (covered with tiny nipple-like projections). Here the phenomenon is
genetically controlled by a single factor, so all the seeds of a given plant are either papillate
or smooth. More common is somatic polymorphism, the production by individual plants of
different seed types, or “morphs.” Somatic polymorphism occurs regularly
in saltbush (Atriplex) and goosefoot (Chenopodium), in which a single plant may produce
both large brown seeds capable of immediate germination and small black ones with some
innate dormancy. Somatic polymorphism may be controlled by the position of the two (or
more) seed types within one inflorescence (flower cluster) or fruit, as in cocklebur, or it may
result from environmental effects, as in Halogeton, in which imposition of long or short days
leads to production of brown or black seeds, respectively. Since the different morphs in
seed (and fruit) polymorphism usually have different dispersal mechanisms and
dormancies, so germination is spread out both in space and in time, the phenomenon can
be seen as an insurance against catastrophe.
Agents Of Dispersal
While some seeds are dispersed independently of the fruits they matured in, others are
dispersed together with the fruit, as is common in many edible fruits, nuts, and cereals.
Such a dispersal unit is referred to as a diaspore. The dispersing agents for seeds and
diaspores are indicated in such terms as anemochory, hydrochory, and zoochory, which
mean dispersal by wind, water, and animals, respectively. Within the zoochorous group,
further differentiation according to the carriers can be made: saurochory, dispersal
by reptiles; ornithochory, by birds; myrmecochory, by ants. Or the manner in which the
seeds or diaspores are carried can be emphasized, distinguishing endozoochory, seeds or
diaspores carried within an animal; epizoochory, seeds or diaspores accidentally carried on
the outside; and synzoochory, seeds or diaspores intentionally carried, mostly in the mouth,
as in birds and ants.
Dispersal by animals

Snails disperse the small seeds of a very few plant species (e.g., Adoxa). Earthworms are
more important as seed dispersers. Many intact fruits and seeds can serve as fish bait,
those of Sonneratia, for example, for the catfish Arius maculatus. Certain Amazon
River fishes react positively to the audible “explosions” of the ripe fruits of Eperua
rubiginosa. Fossil evidence indicates that saurochory is very ancient. The giant
Galapagos tortoise is important for the dispersal of local cacti and tomatoes. The
name alligator apple for Annona glabra refers to its method of dispersal, an example of
saurochory. Many birds and mammals, ranging in size from mice and kangaroo
rats to elephants, eat and disperse seeds and fruits. In the tropics, chiropterochory
(dispersal by large bats such as flying foxes, Pteropus) is particularly important. Fruits
adapted to these animals are relatively large and drab in colour, with large seeds and a
striking (often rank) odour. Such fruits are accessible to bats because of the pagoda-like
structure of the tree canopy, fruit placement on the main trunk, or suspension from long
stalks that hang free of the foliage. Examples include mangoes, guavas, breadfruit, carob,
and several fig species. In South Africa, a desert melon (Cucumis humifructus) participates
in a symbiotic relationship with aardvarks—the animals eat the fruit for its water content and
bury their own dung, which contains the seeds, near their burrows.
Furry terrestrial mammals are the agents most frequently involved in epizoochory, the
inadvertent carrying by animals of dispersal units. Burrlike seeds and fruits, or those
diaspores provided with spines, hooks, claws, bristles, barbs, grapples, and prickles, are
genuine hitchhikers, clinging tenaciously to their carriers. Their functional shape is achieved
in various ways—in cleavers, or bedstraw (Galium aparine), and enchanter’s
nightshade (Circaea lutetiana), the hooks are part of the fruit itself; in
common agrimony (Agrimonia eupatoria), the fruit is covered by a persistent calyx (the
sepals, parts of the flower, which remain attached beyond the usual period) equipped with
hooks; in wood avens (Geum urbanum), the persistent styles have hooked tips. Other
examples are bur marigolds, or beggar’s-ticks (Bidens species); buffalo bur (Solanum
rostratum); burdock (Arctium); Acaena; and many Medicago species. The last-named, with
dispersal units highly resistant to damage from hot water and certain chemicals (dyes),
have achieved wide global distribution through the wool trade.
A somewhat different principle is employed by the so-called trample burrs, said to lodge
themselves between the hooves of large grazing mammals. Examples are mule grab
(Proboscidea) and the African grapple plant (Harpagophytum). In water burrs, such as
those of the water nut Trapa, the spines should probably be considered as anchoring
devices.
Dispersal by birds
Birds, being preening animals, rarely carry burrlike diaspores on their bodies. They do,
however, transport the very sticky (viscid) fruits of Pisonia, a tropical tree of the four-o’clock
family, to distant Pacific islands in this way. Small diaspores, such as those of sedges and
certain grasses, may also be carried in the mud sticking to waterfowl and terrestrial birds.

chesnut-mandibled toucanChestnut-mandibled, or Swainson's, toucan (Ramphastos swainsonii) consuming a nut.©

pablo_hernan/Fotolia

Synzoochory, deliberate carrying of diaspores by animals, is practiced when birds carry

seeds and diaspores in their beaks. The European mistle thrush, Turdus viscivorus,
deposits the viscid seeds of European mistletoe (Viscum album) on potential host plants
when, after a meal of the berries, it whets its bill on branches or simply regurgitates the
seeds. The North American (Phoradendron) and Australian mistletoes (Ameyema) are
dispersed by various birds, and the comparable tropical species of the plant
family Loranthaceae by flowerpeckers (of the bird family Dicaeidae), which have a highly
specialized gizzard that allows seeds to pass through but retains insects. Plants may also
profit from the forgetfulness and sloppy habits of certain nut-eating birds that cache part of
their food but neglect to recover everything or drop units on their way to the hiding place.
Best known in this respect are the nutcrackers (Nucifraga), which feed largely on the “nuts”
of beech, oak, walnut, chestnut, and hazel; the jays (Garrulus), which hide hazelnuts
and acorns; the nuthatches; and the California woodpecker (Balanosphyra), which may
embed literally thousands of acorns, almonds, and pecan nuts in bark fissures or holes
of trees. Secondarily, rodents may aid in dispersal by stealing the embedded diaspores and
burying them. In Germany an average jay may transport about 4,600 acorns per season,
over distances of up to 4 km (2.5 miles). Woodpeckers, nutcrackers, and squirrels are
responsible for a similar dispersal of Pinus cembra in the Alps near the tree line.
Most ornithochores (plants with bird-dispersed seeds) have conspicuous diaspores
attractive to such fruit-eating birds as thrushes, pigeons, barbets (members of the bird
family Capitonidae), toucans, and hornbills (family Bucerotidae), all of which either excrete
or regurgitate the hard embryo-containing part undamaged. Such diaspores have a fleshy,
sweet, or oil-containing edible part; a striking colour (often red or orange); no pronounced
smell; a protection against being eaten prematurely in the form of acids and tannins that are
present only in the green fruit; a protection of the seed against digestion—bitterness,
hardness, or the presence of poisonous compounds; permanent attachment; and, finally,
absence of a hard outer cover. In contrast to bat-dispersed diaspores, they occupy no
special position on the plant. Examples are rose hips, plums, dogwood fruits, barberry,
red currant, mulberry, nutmeg fruits, figs, blackberries, and others. The natural and
abundant occurrence of Euonymus, which is a largely tropical genus, in temperate Europe
and Asia, can be understood only in connection with the activities of birds. Birds also
contributed substantially to the repopulation with plants of the island Krakatoa after the
catastrophic eruption of 1883. Birds have made Lantana (originally American) a pest
in Indonesia and Australia; the same is true of wild plums (Prunus serotina) in parts of
Europe, Rubus species in Brazil and New Zealand, and olives (Olea europaea) in Australia.
Mimicry—the protection-affording imitation of a dangerous or toxic species by an edible,
harmless one—is shown in reverse by certain bird-dispersed “coral seeds” such as those of
many species in the genera Abrus, Ormosia, Rhynchosia, Adenanthera, and Erythrina.
Hard and often shiny red or black and red, many such seeds deceptively suggest the
presence of a fleshy red aril and thus invite the attention of hungry birds.
Dispersal by ants
Mediterranean and North American harvester ants (Messor, Atta, Tetramorium,
and Pheidole) are essentially destructive, storing and fermenting many seeds and eating
them completely. Other ants (Lasius, Myrmica, and Formica species) eat the fleshy, edible
appendage (the fat body or elaiosome) of certain specialized seeds, which they disperse.
Most myrmecochorous plants (species of
violet, primrose, hepatica, cyclamen, anemone, corydalis, Trillium, and bloodroot) belong to
the herbaceous spring flora of northern forests. Tree poppy (Dendromecon), however, is
found in the dry California chaparral; Melica and Centaurea species, in arid Mediterranean
regions. The so-called ant epiphytes of the tropics (i.e., species
of Hoya, Dischidia, Aeschynanthus, and Myrmecodia—plants that live in “ant gardens” on
trees or offer the ants shelter in their own body cavities) constitute a special group of
myrmecochores that provide oil in seed hairs. The ancestral forms of these hairs must have
served in wind dispersal. The primary ant attractant of myrmecochorous seeds is not
necessarily oil; instead, an unsaturated, somewhat volatile fatty acid is suspected in some
cases. The myrmecochorous plant as a whole may also have specific adaptations; for
example, cyclamen brings fruits and seeds within reach of ants by conspicuous coiling
(shortening) of the flower stalk as soon as flowering is over.
Dispersal by wind

seed dispersalA demonstration of how shape and design affect seed speed and dispersal, with single-winged seeds autorotating
and descending at a slower rate than double-winged seeds.Displayed
by permission of The Regents of the University
of California. All rights reserved. (A Britannica Publishing Partner)

In the modern world, wind dispersal (although numerically important) reflects the climatic
and biotic poverty of certain regions; it is essentially a feature of pioneer vegetations. The
flora of the Alps is 60 percent anemochorous; that of the Mediterranean garrigue
(a scrubland region) is 50 percent. By making certain assumptions (e.g., for average wind
velocity and turbulence), the “average limits of dispersal”—that is, the distance that 1
percent of the seeds or diaspores can reach—can be calculated for dispersal units of
various construction and weight. This calculation yields values of 10 km (6 miles)
for dandelion (Taraxacum officinale) and 0.5 km (0.3 mile) for European pine (Pinus
sylvestris). Storms result in higher values—30 km (20 miles) for poplar and 200 km (125
miles) for Senecio congestus.
Too much success in dispersal may be ecologically futile, as exemplified by certain
Florida orchids that arise from windblown West Indian seeds but do not multiply because of
the lack of specific pollinators, usually certain bees or wasps. Anemochorous diaspores can
be subdivided into flyers, dust diaspores, balloons, and plumed or winged diaspores; rollers,
chamaechores or tumbleweeds; and throwers, ballistic anemochores. Dispersal by means
of minute dust diaspores produced in huge quantities is comparable to spore dispersal in
lower plants—a “saturation bombing” is required to find the very limited number of targets,
 or favourable growth habitats, that exist. Not surprisingly, it is practiced mostly by
total parasites, such as broomrapes (in which the finding of the specific host is a problem),
and mycoheterotrophs. The inflated indehiscent pods of Colutea arborea, a steppe plant,
represent balloons capable of limited air travel before they hit the ground and become
windblown tumbleweeds. Winged fruits are most common in trees and shrubs, such
as maple, ash, elm, birch, alder, and dipterocarps (a family of about 600 species of Old
World tropical trees). The one-winged propeller type, as found in maple, is called a samara.
When fruits have several wings on their sides, rotation may result, as in rhubarb and dock
species. Sometimes accessory parts form the wings—for example, the bracts (small green
leaflike structures that grow just below flowers) in Tilia (linden). Seeds with a thin wing
formed by the testa are likewise most common in trees and shrubs, particularly in
climbers—jacaranda, trumpet vine, catalpa, yams, butter-and-eggs. Most famous of these is
the seed with a giant membranaceous wing (15 cm [6 inches] long) of the Javan
cucumber (Alsomitra macrocarpa), a tropical climber.


 
Javan cucumber seedWinged seed of the Javan cucumber (Alsomitra macrocarpa).Scott Zona

Wind dispersal: winged fruits of the silver maple (Acer saccharinum).Thase Daniel

Many fruits form plumes, some derived from persisting and ultimately hairy styles, as
in clematis, avens, and anemones; some from the perianth, as in the sedge family
(Cyperaceae); and some from the pappus, a calyx structure, as in dandelion and Jack-go-
to-bed-at-noon (Tragopogon). Plumed seeds usually have tufts of light, silky hairs at one
end (rarely both ends) of the seeds—e.g., fireweed, milkweeds, dogbane. In woolly fruits
and seeds, the pericarp or the seed coat is covered with cottonlike hairs—
e.g., willow, poplar or cottonwood, kapok, cotton, and balsa. In some cases, the hairs may
serve double duty, in that they function in water dispersal as well as in wind dispersal.
In tumbleweeds, the whole plant or its fruiting portion breaks off and is blown across open
country, scattering seeds as it goes; examples include Russian thistle, pigweed, tumbling
mustard, perhaps rose of Jericho, and “windballs” of the grass Spinifex of Indonesian
shores and Australian deserts. Poppies have a mechanism in which the wind has to swing
the slender fruitstalk back and forth before the seeds are thrown out through pores near the
top of the capsule.

Dispersal by water

Many marine, beach, pond, and swamp plants have waterborne seeds, which are buoyant
by being enclosed in corky fruits or air-containing fruits or both; examples of these plants
include water plantain, yellow flag, sea kale, sea rocket, sea beet, and all species of
Rhizophoraceae, a family of mangrove plants. Sea dispersal of the coconut palm has been
well proved; the fibrous mesocarp of the fruit, a giant drupe, provides buoyancy. Once the
nuts are ashore, the mesocarp also aids in the aboveground germination process by
collecting rainwater; in addition, the endosperm has in its “milk” a provision for seedling
establishment on beaches without much fresh water. A sea rocket species with seeds highly
resistant to seawater is gaining a foothold on volcanic Surtsey Island, south
of Iceland. Purple loosestrife, monkey flower, Aster tripolium, and Juncus species (rushes)
are often transported by water in the seedling stage. Rainwash down mountain slopes may
be important in tropical forests. A “splashcup mechanism,” common
in fungi for spore dispersal, is suggested by the open fruit capsule with exposed small
seeds in the pearlwort (Sagina) and mitrewort (Mitella). Hygrochasy, the opening of fruits in
moist weather, is displayed by species of Mesembryanthemum, Sedum, and other plants of
dry environments.

mangrove propaguleMangrove propagules (Rhizophora species) floating near the Great Barrier Reef.Brian Gratwicke

Self-dispersal

Best known in this category are the active ballists, which forcibly eject their seeds by means
of various mechanisms. In the fruit of the dwarf mistletoe (Arceuthobium) of the western
United States, a very high osmotic pressure (pressure accumulated by movement of water
across cell membranes principally in only one direction) builds up that ultimately leads to a
lateral blasting out of the seeds over distances of up to 15 metres (49 feet) with an initial
velocity of about 95 km (60 miles) per hour. Squirting cucumber (Ecballium elaterium) also
employs an osmotic mechanism. In Scotch broom and gorse, however, drying out of the
already dead tissues in the two valves of the seed pod causes a tendency to warp, which,
on hot summer days, culminates in an explosive and audible separation of these valves,
with violent seed release. Such methods may be coupled with secondary dispersal
mechanisms, mediated by ants in the case of Scotch broom and gorse or by birds and
mammals, to which sticky seeds may adhere, in the case of Arceuthobium and squirting
cucumber. Other active ballists are species of geranium, violet, wood sorrel, witch hazel,
touch-me-not (Impatiens), and acanthus; probable champions are Bauhinia purpurea, with a
distance of 15 metres, and the sandbox tree (Hura crepitans), with 14 metres. Barochory,
the dispersal of seeds and fruits by gravity alone, is demonstrated by the heavy fruits
of horse chestnut.

Plumes on the fruits of mountain mahogany (Cercocarpus) coil and uncoil to drive seeds into the soil, thereby exhibiting self-
dispersal.Dennis Brokaw

Creeping diaspores are found in grasses such as Avena sterilis and Aegilops ovata, the
grains of which are provided with bristles capable of hygroscopic movements (coiling and
flexing in response to changes in moisture). The mericarps (fruit fragments of a schizocarp)
of storksbill (Erodium species), when moistened, bury themselves with a corkscrew motion
by unwinding a multiple-barbed, beak-shaped appendage, which, in the dry state, was
coiled.
Atelechory, the dispersal over a very limited distance only, represents a waste-avoiding
defensive “strategy” that functions in further exploitation of an already occupied favourable
site. This strategy is typical in old, nutrient-impoverished landscapes, such as those of
southwestern Australia. The aim is often achieved by synaptospermy, the sticking together
of several diaspores, which makes them less mobile, as in beet and spinach, and by
geocarpy. Geocarpy is defined as either the production of fruits underground, as in the arum
lilies Stylochiton and Biarum, in which the flowers are already subterranean, or the active
burying of fruits by the mother plant, as in the peanut, Arachis hypogaea. In the American
hog peanut (Amphicarpa bracteata), pods of a special type are buried by the plant and
are cached by squirrels later on. Kenilworth ivy (Cymbalaria), which normally grows on
stone or brick walls, stashes its fruits away in crevices after strikingly extending the flower
stalks. Not surprisingly, geocarpy, like synaptospermy, is most often encountered in desert
plants; however, it also occurs in violet species, in subterranean clover (Trifolium
subterraneum)—even when it grows in France and England—and in begonias (Begonia
hypogaea) of the African rainforest.

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.

Stages of germination of a bean seed.Encyclopædia Britannica, Inc.

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.

Afterripening, stratification, and temperature effects

The most difficult cases of dormancy to overcome are those in which the embryos, although
not underdeveloped, remain dormant even when the seed coats are removed and
conditions are favourable for growth. Germination in these takes place only after a series of
little-understood changes, usually called afterripening, have taken place in the embryo. In
this group are many forest trees and shrubs such as pines, hemlocks, and other conifers;
some flowering woody plants such as dogwood, hawthorn, linden, holly,
and viburnum; fruit trees such as apples, pears, peaches, plums, and cherries; and
flowering herbaceous plants such as iris, Solomon’s seal, and lily of the valley. In some
species, one winter suffices for afterripening. In others, the process is drawn out over
several years, with some germination occurring each year. This can be viewed as an
insurance of the species against flash catastrophes that might completely wipe out certain
year classes.
Many species require moisture and low temperatures; for example, in apples, when the cold
requirement is insufficiently met, abnormal seedlings result. Others (cereals, dogwood)
afterripen during dry storage. The seeds of certain legumes—for example, the seeds of
the tree lupin, the coats of which are extremely hard and impermeable—possess a hilum
with an ingenious valve mechanism that allows water loss in dry air but prevents reuptake of
moisture in humid air. Of great practical importance is stratification, a procedure aimed at
promoting a more uniform and faster germination of cold-requiring, afterripening seeds. In
this procedure, seeds are placed for one to six months, depending on the species, between
layers of sand, sawdust, sphagnum, or peat and kept moist as well as reasonably cold
(usually 0–10 °C [32–50 °F]). A remarkable “double dormancy” has thus been uncovered
in lily of the valley and false Solomon’s seal. Here, two successive cold treatments
separated by a warm period are needed for complete seedling development. The first cold
treatment eliminates the dormancy of the root; the warm period permits its outgrowth; and
the second cold period eliminates epicotyl or leaf dormancy. Thus, almost two years may be
required to obtain the complete plant. The optimal temperature for germination, ranging
from 1 °C (34 °F) for bitterroot to 42 °C (108 °F) for pigweed, may also shift slightly as a
result of stratification.
Many dry seeds are remarkably resistant to extreme temperatures, some even cooled to
that of liquid air (−140 °C or −220 °F). Seeds of Scotch broom and some Medicago species
can be boiled briefly without losing viability. Ecologically, such heat resistance is important
in vegetation types periodically ravaged by fire, such as in the California chaparral, where
the germination of Ceanothus seeds may even be stimulated. The major stimulus after a fire
is a butenolide called karrikin that occurs in smoke. Karrikin is derived from the burning
of cellulose. Also important ecologically is a germination requirement calling for a modest
daily alternation between a higher and a lower temperature. Especially in the desert,
extreme temperature fluctuations are an unavoidable feature of the surface, whereas with
increasing depth these fluctuations are gradually damped out. A requirement for a modest
fluctuation—e.g., from 20 °C (68 °F) at night to 30 °C (86 °F) in the daytime (as displayed
by the grass Oryzopsis miliacea)—practically ensures germination at fair depths. This is
advantageous because a seed germinating in soil has to strike a balance between two
conflicting demands that depend on depth. On one hand, germination in deeper layers is
advantageous because a dependable moisture supply simply is not available near the
surface, but, on the other hand, closeness to the surface is desirable because it allows the
seedling to reach air and light rapidly and become self-supporting.
Light and seed germination

Many seeds are insensitive to light, but in a number of species, germination is stimulated
or inhibited by exposure to continuous or short periods of illumination. So stimulated are
many grasses, lettuce, fireweed, peppergrass (Lepidium), mullein, evening primrose, yellow
dock, loosestrife, and Chinese lantern plant. Corn (maize), the smaller cereals, and many
legumes, such as beans and clover, germinate as well in light as in darkness. Inhibition by
light is found in chive, garlic, and several other species of the lily family, jimsonweed, fennel
flower (Nigella), Phacelia, Nemophila, and pigweed (Amaranthus).

The visible solar spectrum, with prominent Fraunhofer lines representing wavelengths at which light is absorbed by
elements.Encyclopædia Britannica, Inc.

Sometimes, imbibed (wet) seeds that do not germinate at all in darkness may be fully
promoted by only a few seconds or minutes of exposure to white light or to karrikin. The
best-studied case of this type, and one that is a milestone in plant physiology, concerns
seeds of the Grand Rapids variety of lettuce, which is stimulated to germination by red light
(wavelength about 660 nanometres) but inhibited by “far-red” light (wavelength about 730
nanometres). Alternations of the two treatments to almost any extent indicate that the last
treatment received is the decisive one in determining whether the seeds will germinate. This
response involves the phytochrome system, a mechanism that involves a pigment called
phytochrome, which allows green plants to absorb red light. Red
light inhibits stem elongation and lateral root formation but stimulates leaf
expansion, chloroplast development, red flower coloration, and spore germination. Such
stimulation by red light can be reversed by exposure to far-red light.

Ecological role of light

Laboratory experiments and field observations indicate that light is a main controller of
seed dormancy in a wide array of species. The absence of light, for example, was found in
one study to be responsible for the nongermination of seeds of 20 out of 23 weed species
commonly found in arable soil. In regions of shifting sands, seeds of
Russian thistle germinate only when the fruits are uncovered, often after a burial period of
several years. Conversely, the seeds of Calligonum comosum and the melon Citrullus
colocynthis, inhabiting coarse sandy soils in the Negev Desert, are strongly inhibited by
light. The survival value of this response, which restricts germination to buried seeds, lies in
the fact that at the surface fluctuating environmental conditions may rapidly create a very
hostile microenvironment. The seeds of Artemisia monosperma have an absolute light
requirement but respond to extremely low intensities, such as is transmitted by a 2-mm-
(0.08-inch-) thick sand filter. In seeds buried too deeply, germination is prevented. The
responsiveness to light, however, increases with the duration of water imbibition. Even
when full responsiveness to light has been reached, maximal germination occurs only after
several light-exposures are given at intervals. In the field, this combined response
mechanism acts as an integrating (cumulative) rain gauge, because the seeds (as
indicated) become increasingly responsive to light, and thus increasingly germinable, the
longer the sand remains moistened.
Certain Juncus seeds have an absolute light requirement over a wide range of
temperatures; consequently, they do not germinate under dense vegetation or in overly
deep water. (Beneath dense canopies, seed germination is inhibited because the green
leaves above intercept and absorb red light.) In combination with temperature, light (in the
sense of day length) may also restrict germination to the most suitable time of year. In birch,
for example, seeds that have not gone through a cold period after imbibing water remain
dormant after release from the mother plant in the fall and will germinate only when the
days begin to lengthen the next spring.
Stimulators and inhibitors of germination

A number of chemicals (potassium nitrate, thiourea, ethylene chlorhydrin, and karrikin) and
plant hormones (gibberellins and kinetin) have been used experimentally to trigger
germination. Their mode of action is obscure, but it is known that in some instances
thiourea, gibberellin, kinetin, and karrikin can substitute for light.
Natural inhibitors that completely suppress germination (coumarin, parasorbic acid, ferulic
acid, phenols, protoanemonin, transcinnamic acid, alkaloids, essential oils, and the plant
hormone abscisic acid) may be present in the pulp or juice of fruits or in various parts of the
seed. The effect of seed-coat phenols, for example, may be indirect; being highly
oxidizable, they may screen out much-needed oxygen. Ecologically, such inhibitors are
important in at least three ways. Their slow disappearance with time may spread
germination out over several years (a protection against catastrophes). Furthermore, when
leached out by rainwater, they often serve as agents inhibiting the germination of other
competitive plants nearby. Finally, the gradual leaching out of water-soluble inhibitors
serves as an excellent integrating rain gauge. Indeed, it has been shown that the
germination of certain desert plants is not related to moisture as such but to soil water
movement, i.e., to the amount and duration of rain received.
Hans Lambers