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SEED FORMATION, DEVELOPMENT, STRUCTURE AND IMPORTANCE

The purpose of a flower is the production of seed and the purpose of


seed is the production of flower and fruit. Fruit is a ripened ovary
containing seeds. Seed is a ripened ovule containing embryo or unit of
reproduction. A true seed is defined as a fertilized mature ovule consisting
of embryo, stored material and protective coats. Important events involved
in seed formation, development and maturation include
1). Pollination 2). Fertilization 3). Development of the fertilized ovule
by a). cell division, b). accumulation of reserve food material and c).
dehydration.
Floral biology

Flower is a reproductive organ bearing pistil, stamen and usually


sepals and petals. Male part of a flower is androecium consists of anther
sac, anthers and pollen grains. Female part is gynoecium consists of ovary,
style and stigma. The flower is said to be perfect, when they contain both
male and female parts.
A flower with both functional male and female is called as bisexual or
hermaphrodite. Sometimes male or female mature slightly at different
times. If male matures first, it is called as protandry, if female –
protogyny. This nature is called dichogamy – favours cross pollination.
Imperfect flowers have either only male or female part. When both occur

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in a same plant, but separately – monoecious, if they occur in different
plants – dioceious.
Seeds usually developed from the fertilized flower for which pollination
(transfer of pollen from anthers to stigma) is pre-requisite. For successful
fertilization, viable pollen and receptive stigma are two pre-requisites.
The mature anthers dehisce and release pollen grains (haploid
microspores). When pollen grains are transferred from an anther to the
stigma of the same flower, the process is called self-pollination or
autogamy. If they are transferred to the stigma of another flower, cross-
pollination or allogamy is said to have occurred.
Self-pollination occurs in those plants where bisexual flowers achieve
anther dehiscence and stigma receptivity simultaneously. The majority of
angiosperms bear chasmogamous flowers. In some plants, flowers do not
open before pollination, such flowers are called cleistogamous, and this is
the most efficient floral adaptation for promoting self-pollination.
Cross-pollination is ensured in plants which bear unisexual flowers. In
bisexual flowers also self-pollination may be prevented by self-sterility,
dichogamy (maturation of male and female organs at different times),
herkogamy (where the structure of male and female sex organs proves a
barrier to self pollination) and heterostyly (where flowers are of different
types depending on the length of the style and stigma and pollination occurs
only between 2 dissimilar types).
The important self-pollinated crops are wheat, rice, barely, mungbean
and cowpea and cross pollinated are maize, rye, forage legumes and
vegetables like carrot, cauliflower and onion. There is yet another category of
crops called often cross pollinated crops such as cotton and pigeon pea
where there may be 10-40 % cross pollination.
Ovules are developed in the female gametophyte and pollen in the
male gametophyte. Fertilization always takes place in female gametophyte,
therefore, pollen must be transferred from male to female by pollen vector
which may be abiotic including wind (anemophily) and water (hydrophily)
or biotic including insects (entomophily) and bats (cheiropterophily).

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Important steps involved in seed formation
1. Microsporognesis
2. Megasporogenesis
3. Pollen germination
4. Double fertilization
5. Development of embryo and other components
6. Seed maturity
Male and female gametophyte development
Microsporogensis

The process occurs with in the anther which finally results in the
formation of pollen grains. The microspore mother cell undergoes two
nuclear divisions (called meiosis) which result in the formation of four

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haploid functional cells called microspores which upon equal division
become the pollen grains. This is microsporogenesis. The pollen nuclei
further divides mitotically into two each called sperm with ‘n’ number
sperm cells (gametes) viz., tube cell and generative cell. Generative
cell undergoes a division to form two sperm nuclei.
Megasporogenesis
Starts with the enlargement of one cell within a diploid ovule which
have the megaspore mother cell (MMC). MMC undergoes a meiotic
division, giving rise to four megaspores , each containing a haploid
chromosome set. Among the four cells one megaspore survive to give rise
to an embryosac. Whereas other three aborts. The nucleus within the
functional magaspore undergoes three successive divisions to form eight
nuclei. Eight nuclei are arranged as three antipodal cells at the chalazal
end, two central polar nuclei, one egg cell with two synergids at the
micropylar end. Development of eight celled embryosac from the functional
magaspore is called as magagametogenesis.

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Pollen germination
After pollination, the pollen grains reach the stigma and start to
germinate. Pollen germination is dependent upon the receptivity of the
stigma, a condition assumed to be associated with the secretion of the
stigmatic fluid. The surface of the stigma secretes substances which may
provide optimum conditions for pollen germination and subsequent pollen
tube growth.

A long slender pollen tube grows through the style carrying with it, the
tube nucleus and generative nucleus. The generative nucleus soon
divides forming two male gametes, which the tube nucleus gets disorganized
sooner or later. The pollen tube runs down the wall of the ovary and finally it
turns towards the micropyle of the ovule and reaches the embryo sac. This
process is called porogamic fertilization which is the normal method.
Porogamy is the entry of pollen tube through micropyle and chalazogamy
is the entry of pollen tube through chalaza, while entry through integuments
is known as mesogamy. After the pollen tube enters the embryo sac, its tip
dissolves and the male gametes are set free.
Double fertilization
Fertilization involves the union of both sperm nuclei from the pollen
tube with the egg nucleus and the polar nuclei of the embryo sac. One
sperm nucleus unites with the egg nucleus to form the diploid zygote
(embryo) while the other fuses with the polar nuclei or with a single
nucleus, in case the polar nuclei have previously fused, to produce a triploid
nucleus which forms the primary endosperm nucleus. This union of the
two sperm nuclei with the nuclei in the embryo sac is called double
fertilization.

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Failure to set fruit is occasionally due to the fact that double
fertilization did not occur seems probable. Fertilization depends upon the
chromosome constitution of the nuclei involved in the process. In certain
varieties of many species, fruits may develop without fertilization and such
fruits are termed parthenocarpic.

Triple fusion

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Fusion of male gamete with polar nuclei produce triploid nucleus which
is the primary endosperm nucleus. Sometimes there is syngamy or triple
fusion. Syngamy is the fusion of the male gamete with the egg. The time
involved between pollination and fertilization varies a good deal in different
plants. Generally, the time taken is from a few hours to a few days, but in
some cases from a few to several months.
Embryo development
The first division of the zygote is transverse and it results in a small
apical cell and a large basal cell. The apical cell divides vertically forming two
just aposed cells and basal cells undergoes a transverse division forming two
superimpose cells. These result in a T-shaped 4-celled proembryo. The
first superimpose cell divides transversely giving rise to n& n. These two
cells divide further resulting in a row of 3 or 4 cells, forming suspensor.

The second superimpose cell and its derivatives undergo vertical


divisions forming a group of 4 to 6 cells. This group divides by oblique-
periclinal walls forming a set of inner cells and a row of outer cells. The inner
cells form the initials of the root apex and the outer cells from the root cap.
The 2 cells formed as a result of the division of apical cell again divide

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vertically forming quadrants. Each cell of the quadrant divides transversely
and thus can octant containing two tiers of cells are formed.
The cells of the octant undergo vertical division resulting in a globular
proembryo. Periclinal divisions occur in the peripheral cells of the globular
proembryo which delimit an outer layer, the dermatogen. The tier gives rise
to cotyledons and shoot apex and other forms hypocotyle-radicle axis. In
monocotyledons, the apical cell remains undivided and develops into a
haustorial division. The terminal cell of these two by repeated divisions in
different planes fives rise to a single cotyledon. The other cell by a number
of divisions gives rise to stem apex, hypocotyle and root tip.
Endosperm Development
There are 3 types of endosperm development (a) nuclear - where the
endosperm nucleus undergoes several divisions prior to cell wall formation,
e.g., wheat apple, squash, (b) cellular -in which there is no free nuclear
phase, and (c) helobial where the free nuclear division is preceded, and is
followed by cellularization as in some monocots. During the course of seed
development, reserve food materials are accumulated in the endosperm from
the adjacent tissues.
Seed Coat development
The seed coat may be derived from both the integuments or from the
outer integument only; the inner integument may disintegrate. The seed
coat or testa is developed from one or two integuments of the ovules, which
from a covering to protect. (a) the embryo against drying out, (b) the
mechanical injury and (c) attacks by insects, fungi and bacteria. Usually the
outer coat is hard dry and durable (Exine) and the inner (intine) one is thin
and membraneous. Often the two seems to be fused into one layer. The
seed coat and fruit wall may develop appendages or special structures that
help the seed to certain modes of dissemination and protection.
Components of Seed
Seed coat
It is the outer covering of seed and gives protection. It develops from
the 2 integuments of ovule. Outer layer of the seed coat which is smooth and

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rough is known as the testa and is formed from the outer integument. The
inner layer of the seed coat is called the tegmen and is formed from inner
integument.
Embryo
It is the mature ovule consisting of an embryonic plant together with a
store of food, all surrounded by a protective coat, which gives rise to a plant
similar to that of its mother. It is a miniature plant consists of plumule,
radicle and cotyledon. The plumule and radical without the cotyledon is
known as primary axis.
Radicle :Rudimentary root of a plant compressed in the embryo is the
radicle, which forms the primary root of the young seedling. It is enclosed in
a protective cover known as coleorhiza.
Plumule :It is the first terminal bud of the plant compressed in the embryo
and it gives rise to the first vegetative shoot of the plant. It is enclosed in a
protective cover known as coleoptile.
Cotyledon :Cotyledons are the compressed seed leaves. A single cotyledon
(Scutellum) is present in monocots while two cotyledons are present in
dicots, hence they are named as monocots and dicots, respectively. In dicots
they serve as storage tissue and are well developed, while scutellum is a
very tiny structure in monocots.
Endosperm :Endosperm develops from the endosperm nuclei which is
formed by the two polar nuclei and one sperm nuclei. It stores food for the
developing embryo.
Dicot and monocot seeds
Seeds may be broadly classified as dicotyledons and
monocotyledons, depending on the number of cotyledons. Dicotyledons
seeds may be either non-endospermic (exalbuminous) e.g. chickpea, pea and
bean or endospermic (albuminous) e.g., castorbean, fenugreek, etc.,
Monocotyledons seeds are mostly albuminous.

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Monocot seed Dicot seed

Features of a monocot seeds


The seed coat is fused with fruit wall to form pericarp. The endosperm
forms the main bulk of the grain and is the tissue for food storage. It is
separated from the embryo by a definite layer known as epithelium. The
outer most layer of the endosperm is the aleurone layer, which unlike the
rest of the endosperm, is made up of living cells devoid of galactomannan
reserves. This layer secretes alpha-amylase and proteolytic enzymes
which hydrolyse reserves of endosperm.

The embryo is very small and lies in a groove at one end of the
endosperm. It consist of a shield shaped cotyledon (Scutellum) and a short
axis with plumule and radicle protected by root cap. The plumule as a whole
is surrounded by coleptile, a protective sheath, and similarly the radicle
including the root cap is surrounded and protected by coleorrhiza. Scutellum
supplies growing embryo with food material absorbed from endosperm
through epithelium. The initial synthesis of alpha-amylase and certain
proteolases also occurs in scutellum.
Features of a dicot seeds
A typical exalbuminous dicot seed is made up of seed coat and
embryo. The seed coat consists of 2 layers that may be united or free, the
outer layer, which is hard and made of thick walled cells is called testa and
the inner thin membranous layer is called tegument. The seed coat is of

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considerable importance because it is the only protective barrier for the
embryo from the external environment.
The seed coat bears a scar called hilum, marking the point at which
seed is attached to stalk. The funicle or the stalk forms a ridge called raphe
along the margin of the seed. At one end of the hilum, there is a small hole
called micropyle. There is an outgrowth below the hilum in leguminous
seeds, which is called strophiole. Certain other seeds (castorbean, nutmeg)
have outgrowths called arials. Arillar contents may important in attracting
animals, which aid in seed dispersal.
The embryo consists of embyonic axis and 2 fleshy cotyledons.
The axis includes embyonic root (radicle), hypocotyl to which 2
cotyledons are attached and plumule (shoot apex with first true leaves). The
cotyledons of non-endospermic e.g., pea are bulky and account for over 90%
of the mass of the seed.
Cotyledons of epigeal, non-endopermic species become leaf like and
photosynthetic after germination. In endospermic dicot seeds, the
endosperm is bulky and stores food reserves. In these cases, the cotyledons
are small or haustorial in nature. The nucleus of the ovary after fertilization
becomes perisperm. The perisperm in the majority of seeds fails to pass
beyond an incipient stage of development but in a few cases this tissue
becomes the store for food reserves such as coffee.
Seed Growth and Maturation
Wheat and soybean representing monocots and dicots may illustrate
the changes in the pattern of accumulation of reserve materials at different
stages of seed maturation. In wheat, the dry weight of the seed increases
rapidly in about 35 days after anthesis (DAA). The water content of the seed
is maximum between 14 and 21 DAA, and then it declines rapidly. The
amounts of reducing sugar and sucrose are high between 7 and 14 days and
decline rapidly thereafter due to conversion to starch.
The speed of germination is faster in wheat varieties that begin to lose
water early during seed development. The seed is said to have
physiologically matured only when it attains maximum dry weight,

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germinability and vigour. Normally the seed is harvested at field maturity, a
stage when the moisture content is reduced to about 6-10 % in wheat.
A soybean seed attains maximum dry weight between 48 and 54 days
after flowering (DAF). Oil accumulation is less during 12-18 DAF and
maximum between 24 and 42 DAF, after which the rate decreases. The
protein content in the seed is maximum during 12-18 DAF and decreases
subsequently. The initial high percentage of protein may be due to the high
content of non-protein nitrogen, which decreases with seed age. Oil
accumulation picks up only after protein accumulation completes in the seed.
Summary of seed development
1. Translocation of soluble and mobile nutritive substances from the source
tissues.
2. These food reserves are utilized for the condensation and synthesis of
complex storage molecules like starch, protein and lipids takes place
3. These complex molecules start to accumulate in the respective tissues.
Importance of seed structure
11. For identification of cultivars , it can be done based on the morphological,
inheritable physiological and biochemical characters
22. To decide the shelf life potential of seeds -by determining how much
storage organ the seed has
33. To decide about the various post harvest operations namely drying
,threshing , processing ,cleaning, grading to prevent or minimize the
mechanical damage
44. To design post harvest handling equipments.
55. To decide about mechanized sowing.

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