Sexual reproduction

• Sexual reproduction in flowering plant involves the production of male and

female gametes. In a flower, male reproductive structure is the androecium
whereas female reproductive structure is gynoecium.

• It involves the two essential stages: meiosis and fertilization (syngamy).

• Meiosis is required to produce gametes

from diploid structure of sporophyte, while
syngamy is the process of fusion of two
dissimilar gametes: male and female
gametes to form a diploid zygote, thus
restoring a diploid sporophytic phase again.
 Male reproductive structures
Stamen:
• A typical stamen shows two parts: the long and slender
stalk called filament, and terminal generally bilobed
structure called anther. The proximal end of the filament
is attached to the thalamus or the petal of the flower.

• The fertile part of stamen is anther. A typical

angiosperm anther is bilobed. Each lobe has two theca
(dithecous).

• Each lobe contain two microsporangia (pollen sac)

seperated by internal sterile tissue called connective.
Hence, dithecous stamen is tetrasporangiate.
 Development of Microsporangium
• During the development of microsporangium, the anther is seen initially as a
homogeneous mass of meristematic cells, oblong in cross section and
surrounded by a well defined epidermis.

• It then becomes more or less four lobed and in each lobe some hypodermal
cells become more prominent than the others because of their larger size,
more deeply staining cytoplasm and conspicuous nuclei. These cells
constitute the archesporial initials (archesporium. There may be only one
archesporial cell in each of the four lobes as in Boerhaavia.

• The archesporial cells divide by periclinal division (in a plane parallel to

the outer wall of the anther lobe), cutting off primary parietal cells toward
the epidermis and primary sporogenous cells toward the interior of the
anther.
• Then the cells of parietal layer forms 2-5 layers of anther wall by undergoing
a series of divisions, both periclinal as well as anticlinal. The primary
sporogenous cells either directly or after few mitotic divisions functions as
microspore mother cells or pollen mother cells (MMCs or PMCs).

The process of development of anther till the MMCs formation is as follows-

• Archesporial cell- Primary parietal cells and Primary sporogenous cells

• Primary parietal cells- Parietal cells and Endothecium
• Parietal cells- Tapetum and Middle cell layer
• Primary sporogenous cells- Pollen mother cells/Microspore mother cells
Structure of microsporangium (pollen sac)
 Anther and microsporangium
A typical microsporangium is divided into two parts: Anther wall which is
surrounded by four wall layers (the epidermis, endothecium, middle layers and
tapetum) and sporogenous tissue.

1. Anther wall:
i. Epidermis: Outermost single layered thick covering of anther. It is greatly
stretchable and perform the function of protection.

ii. Endothecium: It is single layered, cells are elongated and bear characteristic
fibrous thickening bands of cellulose. This layer attains maximum maximum
growth at the time of dehiscence of pollen. Due to its hygroscopic and fibrous
nature, it helps in dehiscence of anthers at maturity. The cells of endothecium
opposite the partition between two pollen sacs are thin and develop into
stomium.
iii. Middle layers: It may be 1-4 layers or even absent (Lamnaceae, Najadaceae). The
cells of middle layers are ephimerals and degenerate before the pollen mother
cells undergo meiosis; hence it help in early steps of development of pollen
grains.

iv. Tapetum: The innermost wall layer is tapetum which nourishes the developing
pollen grains. The cells of tapetum possess a dense cytoplasm and generally have
more than one nucleus. Tapetum attains maximum development at the stage of
pollen tetrad and degenerate before the dehiscence of anther.

2. Sporogenous tissue:
Inside the tapetum, sporogenous or archesporial cells are present. These cells directly
act as primary micro-sporogenous cells, microspore mother cells or pollen mother
cells. These cells may undergo a few mitosis to add a number of mother cells before
entering into meiosis. Each MMCs undergo meiosis and produces microspore through
microsporogenesis.
 Micro-sporogenesis
• It is the process of formation of microspores from a pollen mother cell
through meiosis.

• A young anther has a group of compactly arranged homogenous cells

called the sporogenous or archesporial cells which occupies the center of
each microsporangium.

• Sporogenous cell on division produce pollen mother cells (PMCs) or

microspore mother cells (MMCs).

• The diploid mother cell produces four functional haploid microspores on

meiosis.
• At the end of meiosis four haploid microspores
are enclosed in a common callose wall.

• The individual spore lacks a wall of its own and

it is a callose partition which separates spores
from each other. Aggregates of four microspores
are called as microspore tetrads. Later on each
spore forms its own wall .

• The arrangement of pollen tetrad may be

tetrahedral (dicot), isobilateral (monocots),
linear (Halophila), decussate (Magnolia) or T
shaped (Butomopsis, Aristolochia).
 Anther dehiscence
• For dehiscence of anther three types of specialized cells are required-
stomium, septum and endothecium.

• The opening through which the pollen grains are discharged from the pollen
sac is called stomium. The septum, that separates the two lobes of an
anther, breaks down at a later stage. The cells of endothecium are thin
walled along the line of dehiscence of each anther lobe.

• On maturity of the anther, a strain is exerted on the stomium due to the loss
of water by the cells of endothecium, with the result the stomium ruptures
and the anther dehisces.

• Mature anther, generally dehisces by means of slit or apical pores.

 Pollen grain
• Pollen grain is haploid, uninucleate, double walled structure.

• It has two layers: outer exine and inner intine which is collectively called
sporoderm.

• Exine is made up of sporopollenin which is one of the most resistant organic

materials known to date. It can withstand high temperatures, strong acids and
alkali. No enzyme can degrade sporopollenin. It has prominent apertures called
germpores.

• The inner wall called intine is a thin and continuous layer made up of cellulose and
pectin.

• The cytoplasm of the pollen grain is surrounded by a plasma membrane.

Mature pollen
 Micro gametogenesis
• The development of male gametophyte from pollen grains is micro
gametogenesis.

• Unicelled pollen grain or microspore represent the first stage of male

gametophytic generation.

• The division of microspore start immediately (tropical plants) or completion

of resting period (cold or temperate plant).

1. Pre- pollination development

• The nucleus of pollen grain mitotically divides into two unequal sized nuclei:
larger vegetative nucleus and smaller generative nucleus.
• Both the nucleus surrounded by cytoplasm and it becomes dense, then followed
by unequal cytokinesis, resulting in two unequal sized cells.

• Initially the generative cell remains at one corner of the spore wall. Within short
time, it gets detached and becomes ellipsoid or fusiform in shape and remains
suspended in the cytoplasm of the vegetative cell (2-celled stage). In majority
of angiosperms, pollen grain shed in this condition.

• This stage of pollen grain is called immature or partially developed male

gametophyte.
2. Post pollination development:
• Further development of pollen grain takes place on the stigma of carpel after
pollination.

• Pollen absorbs moisture and sugar content from stigma, volume of internal contents
increases and exerts pressure on both outer layers.
• As a result, intine comes out through any one germ pore in the form of tube like
structure called pollen tube.

• First of all, vegetative nucleus enter into pollen tube followed by generative nuclei.

• Inside the pollen tube, generative nucleus divides mitotically and form two non-
motile male gametes.

• Pollen grain with one vegetative cell and two male gamete (3- celled structure)
represent the mature male gametophyte of angiosperm.
• Each male gamete has a large nucleus which is surrounded by a thin sheath of cytoplasm.
The tube nucleus may degenerate completely.

• Pollen tube not only carries male gametes but also secretes hormones and absorbs food
from style.
 Female reproductive structures
Carpel
• The gynoecium represent the female reproductive part of the
flower.

• The gynoecium may consists of a single carpel (monocarpellary)

or more than one carpel (multicarpellary).

• When there is more than one carpel, the carpels mat be fused
(syncarpous) or may be free (apocarpous).
• Each carpel has three parts : stigma, style and
ovary.

• Stigma serves as landing platform for pollen

grains. The style is the elongated slender part
beneath the stigma. The basal swollen part of
the carpel is ovary.

• There is ovarian cavity (locules) inside the

ovary, where number of ovules are present.
 Megasporangium/Ovule
• An ovule is a female megasporangium where
the formation of megaspores takes place.

• It is a small structure attached to the placenta

of ovary with a stalk called funicle.

• Hilum represents the point where the body of

the ovule is attached to the funiculus.

• Each ovule has one or two protective layers

called integuments. They are the outer layers
surrounding the ovule that provide protection
to the developing embryo.
• Nucellus – It is a mass of the parenchymatous tissue surrounded by the
integuments from the outside. The nucellus provides nutrition to the
developing embryo.

• The integument encircle the nucellus except at the tip where a small
opening, called micropyle is present. It marks the point where the pollen
tube enters the ovule at the time of fertilization.

• The embryo sac or female gametophyte is located inside the nucellus.

• There is the chalaza opposite to the micropylar end which represents the
basal part of ovule from where the integuments originate.

• Note: The ovule with a single integument is called unitegmic, with two integuments is called
bitegmic and without integument is called ategmic.
 Types of ovule
1. Orthotropous: In this type of ovule, the micropyle is at the distal end and the
micropyle, the funicle and the chalaza lie in one straight vertical line. Examples:
Piperaceae, Polygonaceae

2. Anatropous: The body of the ovule becomes completely inverted so that the
micropyle and funiculus come to lie very close to each other. This is the common
type of ovules found in dicots and monocots.

3. Hemianatropous: In this, the body of the ovule is placed transversely and at right
angles to the funicle. Example: Primulaceae.

4. Campylotropous: The body of the ovule at the micropylar end is curved and more
or less bean shaped. The embryo sac is slightly curved. All the three, hilum,
micropyle and chalaza are adjacent to one another, with the micropyle oriented
towards the placenta. Example: Leguminosae
5. Amphitropous: The distance
between hilum and chalaza is less.
The curvature of the ovule leads to
horse-shoe shaped nucellus. Example:
some Alismataceae.

6. Circinotropous: Funiculus is very

long and surrounds the ovule.
Example: Cactaceae
 Megasporogenesis
• The process of forming megaspores from megaspore mother cell is
known as megasporogenesis.

• Megaspores are produced in ovules. A single cell in each ovule ie.,

hypodermal cell in the nucellus (at the micropylar end) differentiates and
functions as the archesporium (archesporial cell).

• It is distinguishable from the other cells as it becomes more prominent

than its surrounding cells due to its large size, dense cytoplasm and large
nucleus.

• Ovule can be categorized into two types (tenuinucellate and

crassinucellate), on the basis of position of sporogenous cell.
• In tenuinucellate type of ovule, the archesporial cell directly functions as
megaspore mother cell (MMC) and in crassinucellate type of ovule the
archesporial cell do not directly behave as MMC and instead of that it
divides periclinally into two cells.

• An outer primary parietal cell (towards epidermis) and an inner primary

sporogenous cell. Now this primary sporogenous cell functions as the
megaspore mother cell.

• The megaspore mother cell undergoes meiosis and produces a row of four
haploid megaspores (linear tetrad).
• In the linear tetrad the lowermost megaspore (the chalazal megaspore)
enlarges and becomes functional. Rest three megaspores of tetrad do not
participate in the formation of female gametophyte and degenerate.
 Mega gametogenesis
• Megaspore (n) is the first cell of the female gametophyte. The functional mega-
spore becomes enlarged and thus forms the female gametophyte i.e., the embryo
sac. This method of embryo sac formation from a single megaspore is termed as
monosporic development.

• Initially, the embryo sac is uninucleate and with further growth its nucleus divides
by three successive divisions and forms eight nuclei.

• Out of eight nuclei, initially four remain towards the micropyle end and the other
four towards the chalazal end.

• Six of the eight nuclei are surrounded by the cell walls and organized into the
cells, the remaining two nuclei (polar nuclei) from each pole moves towards
centre below the egg apparatus in the large central cell.
• The three nuclei of the micropylar end form the egg apparatus and the rest
three at the chalazal end are called antipodal cells.

• In the egg apparatus, each nucleus is surrounded by viscous mass of cyto-

plasm without any wall, of which the middle one is the largest and called
egg, ovum or oosphere and the rest two (one on each side of the egg) are
the synergids or helping cells.

• The antipodal cells have viscous mass of cytoplasm, covered by cellulosic

wall.

• Thus, a typical angiosperm embryo sac at maturity is 7-celled through it is

an eight-nucleate embryo.
Different stages of development of female gametophyte of angiosperm
Pollination
• The process of transfer of pollen grains from anther (microsporpophyll) to
stigma (receptive region of megasporophyll) is called as pollination.

• Pollination starts from the release of pollen grains from anther and ends
when it reaches to the stigma.

• Pollination is successful when fertilization results.

• The pollination may takes place inside a flower (autogamy or self-

pollination) or in between two flowers (allogamy) of same plant
(geitonogamy) or of different plants (xenogamy) or even in between two
species (hybridization).
 Self -pollination
• The process of transfer of pollen grains from anther to stigma within a
flower / of a same flower. It is referred to as the primary type of
pollination as it includes a single flower.

• Self-pollination occurs when pollen grains fall directly from anther into
the stigma of the flower.

• This process is quite simple and fast, which leads to a reduction in

genetic diversity as the sperm and egg cells of the flower share some
genetic information.
Self-pollination
 Contrivances of self-pollination
1. Bisexuality or uniciliny: For self-pollination, the flower should be bisexual
and perfect. Unisexual flower require pollen grains of another flower
which is cross pollination.

2. Homogamy: The condition in which anther and stigma of a flower mature

at same time is homogamy. Eg. Pea

3. Chasmogamy: Certain plant produces several adaptive characters for ease

of self-pollination.
• In Vinca rosea, Convolvulus and Gardenia, the matured anther borne
near the mouth of corolla tube and stigma.
 • In Mirabilis, anther bend over the stigma; in potato, style bends over the
anthers; while in sunflower stigma curl back to receive pollen.
• In Lilac, stigma lies exactly below the anther. These all conditions favour
self-pollination.

4. Cleistogamy: Usually angiospermic flower opens at maturity

(chasmogamous) but several flower don’t open even at maturity
(cleistogamous). The bisexual cleistogamous flower produce fruits after self
pollination. Eg. Viola, Oxalis, Drosera. The cleistogamy flowers found under
soil produce geocarpic fruits (groundnut).

5. Bud pollination: Before opening of a bud, the pollination take place as in

most cultivated plants that are called bud pollination. Eg. Wheat, tobacco,
jute, rice, etc.
 Advantages
1. Self- pollination ensures that recessive characters are eliminated.

2. The wastage of the pollen grain is very less compared to cross-pollination.

3. In the process of self- pollination, the purity of the race is maintained, as

there is no diversity in the genes.

4. In self- pollination, there is no involvement of external factors like wind,

water, and other pollinating agents.

5. Self-pollination ensures that even a smaller quantity of produced pollen

grains from plants have a good success rate in pollination.
 Disadvantages

The major disadvantage of Self- pollination is there is no mixing up of

genes.
Due to which:
• The vigor and vitality of the race are reduced.
• The immunity to diseases is reduced in the resultant offspring.
Cross pollination (Allogamy)
• The transfer of pollen grains from anther to the stigma of another
flower is cross-pollination.

• It is possible only in the presence of certain vectors or agents such as,

insects, birds, air, water, wind, humans, etc.

• It may take place between two genetically similar flowers, i.e., of

same plant (geitonogamy) or in between two genetically different
flowers i.e., of two plants (xenogamy) or in between two genetically
different plants (hybridization).
 Contrivances or factors of cross pollination

1. Dicliny or unisexuality: The flower, which bears either carpel or stamen,

never both, favors cross pollination. The unisexual flower may be found
in monoecious or dioecious plants.

a) Monoecious: Maize, Cucurbits, Banana, Mango, Cucumber, etc.

b) Dioecious: Canabis, Spinach, Salix, Piper longum, etc.

2. Dichogamy or heterogamy: The condition in which male sex organs

anther and female sex organ pistil mature at two different times is called
dichogamy. It may be protoandry or protogyny.

a) Protoandry: Androecium matures earlier than stigma. Eg.Sunflower, Lady’s finger

b) Protogyny: Gynoecium matures earlier than stamen. Eg. Brassica, Rosa, Mirabilis,
etc.
3. Self- sterility or self-incompatability or genetical barrier: The pollen
grains of certain bisexual flowers are incapable of growing over the
stigma of same flower. They have multiple alleles, which can prevent
self-pollination. Eg. Tobacco, potato, etc.

4. Heretrostyly and heteroanthy: The flowers of several plants have two or

three morphological types of styles and stamens. It may be diheterostyly
or triheterostyly.

a) Diheterostyly flower has two types of style, one pin eyed (long style
with short stamens) and another thrum eyed (short style with long
stamens). Eg. Primula, Jasmine.

b) Triheterostyly flower has three types of styles (long, medium or short)

and stamens and pollination in between same height stamens and
styles of different flowers. Eg. Lythrum, Oxalis, etc.
5. Herkogamy: It is a mechanical barrier as Caryophyllaceous flower, in
which at maturity, the stigma projects beyond the reach of stamen.

• In Clerodendron and Gloriosa, the mature stigma and anthers occur in

different positions and pollens dehiscence at a distance.

• In Calotropis, Orchidaceae and Asclepediaceae, pollen grains occur in

sacs called pollinia. Two pollinia are attached ro common sticky
corpusculum to form a translator, which can be lifted by insects only.

• In Salvia, there is turn pipe mechanism (lever mechanism) to shed

pollen grains at the back of visiting insects.
6. Heteromorphism: Certain plants may produce flowers of different forms.
Eg. Primula.

7. Male sterility: Certain plants may produce bisexual flowers but the
anther become sterile at the time of pollination.

8. Pollen prepotency: The process of quicker growing of pollen grain of

another plant than that of same plant is called pollen prepotency. Eg.
Apple.
Advantages Of Cross-pollination

• The produced seeds are good in vigor and vitality.

• All unisexual plants can reproduce through the process of Cross-
pollination.
• The recessive characters in the lineage are eliminated as a result of
genetic recombination.
• This process improves the immunity of the offsprings towards the
diseases and other environmental factors.
• Cross-pollination introduces new genes into a sequence of
species and this is mainly due to the fertilization between
genetically different gametes.
 Disadvantages
• In this process, there is a great wastage of pollen grains.

• Due to genetic recombination during meiosis, there are

chances of eliminations of good qualities and additions of
unwanted characteristics in offspring.
 Agents for pollination
Pollination process can occur by different agencies which can be classified
into two categories:
 Anemophily or anemogamy:
• Here pollinating agent is wind e.g. in most cereals, poplar, willow, alder, elm, oak, beech,
Urtica.

• The wind-pollinated flowers comprise light-coloured

petals without a pleasant strong smell.
• The produced pollen grains are smaller and lighter in
weight, which can be carried by the wind easily.
• Stigma is feathery or sticky and found hanging out of
petals. The stamens are long and visible out of the petals.
• The anthers are often seen being supported outside the
flower.
• The filaments found in these flowers are slender and
long.
• These flowers do not produce nectars.
 Hydrophily or Hydrogamy
• Here pollinating agent is water e.g. aquatic plants.

• Flowers are light, small and inconspicuous.

• Pollen grains are light protected by mucilage covering and therefore unwettable.

• Female flowers reach the surface of water. Stigma

long, feathery and sticky.

• They produce a large number of pollen grains as most

of them get lost in the water flow and very few are
able to reach the stigma.

• Male flowers are also released on the surface of water

so that they may be carried through water currents to
the stigmas of female flowers to effect pollination at
the water surface as it Vallisneria.
 Entomophily
• Pollinating agents are insects as in Salvia, Ficus, orchids etc.

• The insect-pollinated flowers comprise brightly coloured petals with a pleasant strong smell.

• In insect-pollinated flowers, the produced pollen grains are larger in size, sticky and spiny,
which helps the insect to carry the pollen grains.

• Stigma is small and is situated deep inside the

petals.
• Stamens may be small and hidden inside petals.

• The anthers are found deep inside the flower.

• The filaments found in these flowers are strong
and short.
• These flowers produce a lot of nectars.
 Ornithophily
• Pollinating agents are birds as in Bignonia, silk cotton etc.

• Flowers are large and showy.

• Tubular, cup or urn to ease the entry of beaks

of different birds.
• Produce considerably a large amount of nectar.

• The flowers have attractive and bright colors

such as yellow, red and orange. These bright
colors have a larger wavelength and are visible
even at a longer distance for the birds.

• Flowers are usually fragrant.

 Cheiropterophily:
• Pollinating agents are bats as in Bauhinia megalandra, Eperua falcata etc.
• The flowers are usually large, wide-mouthed to accommodate the head of the bat and
white or pale-colored so that they can be distinguished from their dark surroundings
at night.

• Flowers bloom at night.

• The flowers have a strong, fruity, or musky

fragrance and produce large amounts of nectar.

• As the bats seek the nectar, their faces and heads

become covered with pollen, which is then
transferred to the next flower.
 Hybridization

• It is the process of crossbreeding

between genetically dissimilar
parents to produce a hybrid.

• It is an artificial method carried

out by humans.
Fertilization
• The fusion of two dissimilar gametes (male gamete and female gamete) is
fertilization. It is also called syngamy or fecundation.

• The fertilization leads to the formation of diploid zygote or oospore.

• After pollination, pollen grains reach at stigma. The intine of each pollen
grain grows out into a tube (pollen tube) through thin, weak germ pore of
exine. The process of fusion of gametes by means of pollen tube is
siphonogamy.

• In gymnosperm, pollen grain directly reach at mouth of ovules due to

absence of stigma, style and ovary, hence siphonogamy is absent.
• Usually the pollen grain has two cells at pollinating stage: generative cell
and vegetative cell. The generative nucleus soon divides and gives identical
two male gametes, while tube nucleus soon or later disorganizes.

• The pollen tube shows chemotactic

movement ie., movement of growth due to
chemical (glucose or malic acid).

• The pollen tube is guided and nourished

by other tissues in ovary called obturator
(outgrowth of placenta or funicle that
surrounds nucellus.
 Entry of pollen grain to ovules
After arriving in the ovary, the pollen tube finds its way into the ovule. The
pollen tube may enter into the ovule via three routes.
i. through the micropyle
ii. through the chalazal end
iii. through the integument

On that basis of modes of entry of pollen tube into the ovule, three terms are
given as follows:
1. Porogamy: When the pollen tube enters the ovule through the micropyle,
the condition is known as porogamy. This is the most common mode of
pollen tube entry into the ovule and so the most common type of
fertilization.
2. Chalazogamy: When the pollen tube enters the ovule through the chalazal end, the
condition is known as chalazogamy. This type of pollen tube entry into the ovule
and so the type of fertilization is observed in Casuarina, Betula and Juglans regia.
The chalagogamy was first reported by Treub (1891) in Casuarina.

3. Mesogamy: When the pollen tube enters the ovule through the integument or
through the funiculus, the condition is known as mesogamy. This type of pollen tube
entry into the ovule and so the type of fertilization is observed in Cucurbita (through
the integument), and Pistacia (through the funiculus).

Therefore depending on the place of pollen tube entry into the ovule, fertilization may
also be called of three types: Porogamous, Chalazogamous and Mesogamous.
Discharge of male gametes from pollen tube
• After reaching the embryo sac the pollen tube burst at its tip and deliver the (two)
male gametes. Just prior to bursting of pollen tube the tube nucleus disorganizes.
• Immediately after releasing, the male gametes show amoeboid movement and one
male gamete moves toward the egg and other one move to the polar nuclei.

Syngamy- fusion of gametes

• As the one of the male gametes reached the egg, it fuses with it. As a result of this
fusion diploid zygote/oospore (2n) forms. The fusion of male and female gametes
is known as fertilization. This is also known as syngamy.

• Since two male gametes are released by the pollen tube, one male gamete fused
with the egg (syngamy) and the other male gamete with the polar nuclei (triple
fusion)
Triple fusion
The other male gamete fuses with the two polar nuclei (or secondary nucleus, if the
two have already fused) and so forms triple fusion nucleus (3n), called primary
endosperm nucleus.

Double fertilization
• Thus, in an embryo sac two sexual fusions occur; one is syngamy (i.e. fusion of
one male gamete with the egg) and another is triple fusion (i.e. fusion of other
male gamete with the polar nuclei or secondary nucleus), and therefore, the
phenomenon is known as double fertilization.

• As a result of first fertilization the zygote or oospore cell is formed which is the
mother cell of the embryo and is a diploid cell containing 2n complement of the
chromosomes. The nucleus of the triple fusion product (primary endosperm
nucleus) is triploid or 3n. This is the first nucleus of the endosperm.
• Double fertilization is a very unique phenomenon in Angiosperms and
discovered for the first time by S.G. Nawaschin (1898) in Lilium and
Fritillaria species.
 Post Fertilization Developments

• After fertilization, development of the embryo and the endosperm within

the ovule goes side by side.

• The oospore (zygote), formed as a result of fusion of one male gamete with
the egg, develops into the embryo while the primary endosperm nucleus-
product of triple fusion, develops the endosperm.

• The other nuclei or cells within the embryo sac (synergids, antipodal cells)
disorganize sooner or later.
 Development of the Endosperm
• The primary endosperm undergoes a series of divisions and ultimately forms endosperm.

• The Angiospermic endosperm is a triploid (3n). It is formed either by the fusion of one
haploid male gamete and one diploid secondary nucleus (fusion product of two haploid
polar nuclei) or by the fusion of three haploid nuclei (one male gamete belongs to male
gametophyte and two polar nuclei belongs to the female gametophyte).

• It is therefore distinct from the endosperm of heterosporous: Pteridophytes and

Gymnosperms where the endosperm is a simple haploid (n) tissue of the gametophyte not
involving any triple fusion like in Angiosperms. Endosperm is a highly nutritive tissue
which provides nourishment to the developing embryo.

• In Orchidaceae and Podostemonaceae, the product of double fertilization soon

disintegrates and endosperm development is completely suppressed.
Depending upon mode of development three types of endosperm has been
recognized:

1. Nuclear endosperm
2. Cellular endosperm
3. Helobial endosperm

Of these nuclear endosperm is the most common type which occurs in about
56% families of Angiosperms. It is followed by cellular endosperm (reported
in 25% families of Angiosperms) and then by helobial endosperm (reported in
19% families of Angiosperms).
 Nuclear Endosperm
• In this type of endosperm the division of primary endosperm nucleus and
number of subsequent nuclear divisions are not accompanied by wall
formation and the nuclei thus produced remain free in the cytoplasm of the
embryo sac.

• They remain in the peripheral layer of the cytoplasm surrounding a large

central vacuole. Wall formation occurs at later stage around nuclei. The wall
formation is mostly centripetal i.e. from the periphery towards the centre
and usually begins from the basal periphery e.g. Arachis hypogea.

• In some cases, the central vacuole may not be filled up even in the mature
seed. This is seen in the palms.
• Cocus nucifera is the classical example of this type of nuclear endosperm. The primary
endosperm nucleus undergoes a number of free nuclear divisions. Then the embryo sac
gets filled with a clear fluid (watery liquid endosperm) in which numerous nuclei float.
It is known as liquid syncytium. Gradually nuclei start settling at the periphery with the
beginning of peripheral cell wall formation. This forms the coconut meat.

• In mature coconuts the liquid endosperm becomes milky. The watery endosperm of
coconut contains growth promoting „coconut milk factor‟ and that is why it is used as a
nutrient medium in plant tissue culture experiments. Nuclear endosperm is commonly
found in polypetalous dicotyledons .
 Cellular Endosperm
• In this type of endosperm, division of the primary endosperm nucleus is
immediately followed by wall formation so that the endosperm is
cellular from the beginning. The first wall is laid down transversely but
the subsequent divisions are irregular. Adoxa, Peperomia, Villarsia etc.
are some common examples.
 Helobial Endosperm
• Among members of Helobiales (e.g. Vallisneria, Eremurus, Limnophyton etc.)
there is type of endosperm the development which is intermediate between the
nuclear and the cellular type.

• Here, the first division of the primary endosperm nucleus is accompanied by

the formation of transverse wall. This divides the embryo sac unequally into
two compartments - a small chalazal chamber and a large micropylar chamber.

• This step is followed by free nuclear division in both the chambers but there
are relatively more free nuclear divisions in micropylar chamber in
comparison to chalzal one. The chalazal chamber often degenerates.
• The free nuclear divisions in the micropylar chamber are followed by
wall formation and thus a cellular endosperm tissue is formed and
usually found in the members of the order Helobiales.
Seeds can also be categorized into two categories.
1. Non-endospermic seeds
2. Endospermic seeds

1. Non- endospermic seeds (ex-albuminous seeds)

• In plants where the entire endosperm consumed or utilized in the nutrition of the
developing embryo, the mature seeds thus formed are without endosperm. Such
seeds are termed as non-food material in cotyledons.

2. Endospermic seeds (albuminous seeds)

• In plants where the seeds retain endosperm even at maturity and do not consumed
or utilized the endosperm completely in the nutrition of the developing embryo.
Such seeds are said to be endospermic seeds. Example are seeds of coconut, castor
etc. The endosperm present in the seed is utilized after germination in the
establishment of young seedlings.
 Development of the embryo
• After fertilization, a series of changes occurs in the ovule and finally seed is
formed. Side by side with the development of the endosperm, the oospore
(zygote, the fertilized egg) is developing the embryo after a period of rest.

• The process of development of mature embryo from diploid oospore is called

embryogenesis.

• Both dicotyledons and monocotyledons begin embryo development in the

same way but there is considerable difference in later differentiation.

• The dicotyledonous embryo as the name reflects, has two cotyledons attached
laterally to an embryonical axis, whereas in the monocotyledonous embryo,
the embryonical axis has a single cotyledon at its apex.
• Due to this organographic difference, it is very easy to distinguish the two
types of embryo but there is no fundamental difference in their early stage of
development. The development is very similar till the globular stage.

• In all Angiosperms, the embryogenesis starts with the division in oospore and
it divides to develop a two-celled proembryo by forming a transverse wall.

• The cell near the micropyle is termed the basal cell and the cell facing towards
the centre of the embryo sac is called the terminal cell.

• The basal cell forms the suspensor and may or may not contribute in rest
activities so sometimes called as suspensor cell, whereas terminal cell is
responsible for further development of embryo so called embryo cell.
 Types of embryo development
On the basis of plane of division of the terminal cell (also known as apical or embryo
cell) in the 2-celled proembryo and the contribution of the basal cell and terminal
cells in the formation of embryo proper, six types of embryogeny (embryo
development) have been reported by Johansen (1950) among the Angiosperms.

1. Onagrad or Crucifer type (e.g. Annonaceae, Brassicaceae, Onagraceae,

Pedaliaceae, Ranunculaceae, Scrophulariaceae).
2. Asterad type (e.g. Asteraceae, Balsamianceae, Violaceae, Vitaceae).
3. Solanad type (e.g. Campanulaceae, Linaceae, Solanaceae, Theaceae).
4. Caryophyllad type (e.g. Caryophyllaceae, Crassulaceae,Haloragaceae).
5. Chenopodiad type (e.g. Boraginaceae, Chenopodiaceae).
6. Piperad type (e.g. Loranthaceae, Piperaceae).
 Development of dicotyledonous embryo

• Hanstein (1868) gave the typical type of dicot embryogenesis in Capsella

bursa-pastoris (shepherd's purse) of Brassicaceae. Later Mahshwori (1950)
gave five different types of embryogenesis, out of them Onagrad or
Crucifer type is most common.

• The ovule is campylotropous so that the embryo sac and the later developed
endosperm as well as embryo are horseshoe-shaped.

• Zygote (oospore) divides transversely. To give two unequal cells: larger

basal cell at the micropylar end is called suspensor cell. The smaller one,
away from it termed as terminal cell or embryo cell. As a result of this a
two-celled proembryo is formed.
• The suspensor cell divides transversely a few times to produce a
filamentous suspensor of 6-10 cells. The suspensor helps in pushing the
embryo in the endosperm.

• The first cell of the suspensor (towards micropyle) becomes swollen and
called haustorium or vesicular cell.

• The last cell of suspensor (towards embryo cell) is known as hypophysis. It

forms radicle and root cap.

• The embryo cell undergoes two vertical divisions and one transverse
division to form quadrant and then octant stage. In octant, eight cells
arranged in two tiers- epibasal (terminal) and hypobasal (near the
suspensor).
• The epibasal cells eventually form the two cotyledons and the plumule.

• The hypobasal cells produce the hypocotyl. For this the octant embryo
undergoes periclinal divisions producing outer layer of protoderm,
procambium and ground meristem.

• Protoderm forms epidermis, procambium gives rise to steal or vascular

strand and ground meristem produces cortex and pith.

• It is initially globular but with the growth of cotyledons it becomes heart-

shaped and then assumes the typical shape, e.g., Capsella bursa-pastoris.
 Structure of Dicot Embryo
• The mature embryo consists of an embryonal
axis having two cotyledons.

• Embryonal axis above the level of cotyledons

forms the plumule (epicotyl) and below the
cotyledons, the radical (hypocotyl).

• Upon germination the plumule forms the

shoot and the radical gives rise the root
system. The reserve food material in the
cotyledons is used in the establishment of
young seedlings.
 Development of monocotyledonous embryo
• There is no essential difference between the embryogeny of monocotyledons
and that of dicotyledons but as a single cotyledon develops instead of two
from the embryo in monocotyledons, there is some difference in later stages.

• We are taking an example of Luzula forsteri of Juncaceae for describing the

development of monocotyledonous embryo. Here the development of
embryo is also Onagrad or Crucifer type.

• The early development of dicot and monocot embryos are similar upto
octant stage. Later on differentiation starts.
1. The zygote or oospore elongates and then divides transversely to form basal and
terminal cells.
2. The basal cell (towards micropylar end) produces a large swollen, vesicular
suspensor cell. It may function as haustorium.

3. The terminal cell divides by another transverse wall to form two cells.
4. The top cell after a series of divisions forms plumule and a single cotyledon.
5. Cotyledon called scutellum, grows rapidly and pushes the terminal plumule to one
side. The plumule comes to lie in a depression.

6. The middle cell, after many divisions forms hypocotyl and radicle. It also adds a few
cells to the suspensor.

7. In some cereals, both plumule and radicle get covered by sheaths developed from
scutellum called coleoptile and coleorhiza respectively.
 Structure of Monocot Embryo
• The embryos of monocotyledons have only one cotyledon.

• In grass family (Poaceae), this cotyledon is called

scutellum. It is situated towards lateral side of embryonal
axis. This axis at its lower end has radicle and root cap
enclosed in a sheath called coleorhiza.

• The part of axis above the level of attachment of scutellum

is called epicotyl.

• It has shoot apex and few leaf primordia enclosed in a

hollow foliar structure called coleoptile. Epiblast represents
rudiments of second cotyledon.