Seed

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 of various plants. Row 1: poppy, red pepper,
strawberry, apple tree, blackberry, rice, carum, Row
2: mustard, eggplant, physalis, grapes, raspberries,
red rice, patchouli, Row 3: figs, lycium barbarum,
beets, blueberries, golden kiwifruit, rosehip, basil,
Row 4: pink pepper, tomato, radish, carrot,
matthiola, dill, coriander, Row 5: black pepper, white
cabbage, napa cabbage, seabuckthorn, parsley,
dandelion, capsella bursa-pastoris, Row 6:
cauliflower, radish, kiwifruit, grenadilla, passion fruit,
melissa, tagetes erecta.

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
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

Stages of seed development:

I Zygote
II Proembryo
III Globular
IV Heart
V Torpedo
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 …
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 …

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 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
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 tegmen 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 pericarp.) 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 testae are homologous from
one species to the next. The funiculus
abscisses (detaches at fixed point –
abscission zone), the scar forming an
oval depression, the hilum. Anatropous
ovules have a portion of the funiculus
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. Gossypium 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)

Gymnosperms …

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

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

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

seed coat and embryo
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 …

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. Scutellum B.
Cotyledon C. Hilum D. Plumule E. Radicle F.
Endosperm

Comparison of monocotyledons and dicotyledons

Embryo …

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 …

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 …

Seed coat of pomegranate

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 …

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 autumn
all 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
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 …

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 …

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) …

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) …

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) …

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 …
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.
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 …

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.

Far red light can prevent germination.[36]

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.[37]

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.[38] 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 …

During seed dormancy, often associated

with unpredictable and stressful
environments, DNA damage
accumulates as the seeds age.[39][40][41]
In rye seeds, the reduction of DNA
integrity due to damage is associated
with loss of seed viability during
storage.[39] Upon germination, seeds of
Vicia faba undergo DNA repair.[40] 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.[42] Also, in
Arabidopsis seeds, the activities of the
DNA repair enzymes Poly ADP ribose
polymerases (PARP) are likely needed for
successful germination.[43] 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 …

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.[44] 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.[45]

Sterile seeds …
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 …

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.

True to the seed …

Also called growing true, refers to plants
whose seed will yield the same type of
plant as the original plant. Open
pollinated plants, which include
heirlooms, will almost always grow true
to seed if another variety does not cross-
pollinate them.

Economic importance

Phaseolus vulgaris (common bean or green bean)

seeds are diverse in size, shape, and color.

Seed market …
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.[46]

Edible seeds …

Many seeds are edible and the majority

of human calories comes from seeds,[47]
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.[48] 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 …

While some seeds are edible, others are

harmful, poisonous or deadly.[49] 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.[50]

A deadly poison, ricin, comes from seeds

of the castor bean. Reported lethal doses
are anywhere from two to eight
seeds,[51][52] though only a few deaths
have been reported when castor beans
have been ingested by animals.[53]

In addition, seeds containing

amygdalin – apple, apricot, bitter
almond,[54] peach, plum, cherry, quince,
and others – when consumed in
sufficient amounts, may cause cyanide
poisoning.[54][55] 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.[51][56] 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.[57]

Other uses …

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

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.[58]
(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.[59][60])
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.[61]
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.[62]
 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.[63]

In religion
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.[64]

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?[65]

See also
Biological dispersal
Carpology
Genetically modified crops
List of world's largest seeds
Recalcitrant seed
Seed company
Seed enhancement
Seed library
 Seed orchard
Seed paper
Seed saving
Seed testing
Seed trap
Seedbed
Soil seed bank
Selective embryo abortion

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External links

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Royal Holloway, University of London:
The Seed Biology Place
The Millennium Seed Bank Project
Kew Garden's ambitious preservation
project
The Svalbard Global Seed Vault – a
backup facility for the world's seed
banks
Plant Physiology online: Types of Seed
Dormancy and the Roles of
Environmental Factors
Canadian Grain Commission:Seed
characters used in the identification of
small oilseeds and weed seeds
The Seed Site : collecting, storing,
sowing, germinating, and exchanging
 seeds, with pictures of seeds,
seedpods and seedlings.
Plant Fix: Check out various plant
seeds and learn more information
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