02

SEXUAL REPRODUCTION IN FLOWERING PLANTS

KEYNOTES 2.0
By: Dr. Anand Mani

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 All flowering plants (angiosperms) show sexual reproduction. Flowers are the
sites of sexual reproduction

PRE FERTILISATION: STRUCTURES & EVENTS

Several hormonal and structural changes result in differentiation and development of the floral primordium.

Inflorescences bear the floral buds and then the flowers.

ANDROECIUM (STAMEN)
It consists of a whorl of stamens representing the male reproductive organ. Their number and length are
variable in different species.

A stamen has 2 parts:

Filament: Long and Anther: Terminal and typically

slender stalk. Its proximal bilobed. Each lobe has 2 thecae
end is attached to the (dithecous), often a longitudinal
thalamus or the petal of groove runs lengthwise
the flower. [NEET-I 2016] separating the theca.

Transverse section of anther:

The anther is a tetragonal structure consisting of four
located at the corners.

Each lobe consists of two microsporangia. Hence, the

dithecous anthers are tetrasporangiate.

The microsporangia develop to pollen sacs. They extend

longitudinally all through the length of an anther and
are packed with pollen grains.
Structure of a microsporangium
A typical microsporangium is near circular in outline.

It is surrounded by four wall layers– the epidermis,

endothecium, middle layers & tapeturn

The outer 3 layers give protection and help in dehiscence

of anther to release the pollen.

The tapeturn (innermost layer) nourishes the developing

pollen grains. [NEET 2013]

Cells of the tapetum contain dense cytoplasm and

generally have more than one nucleus.

When the anther is young, a group of compactly

arranged homogenous cells (sporogenous tissue) occupies
the centre of each microsporangium.

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Microsporogenesis:

As the anther develops, each cell of sporogenous tissue undergo meiotic divisions to form
microspore tetrads (microspores are arranged in a cluster of four cells). Each one is a
potential pollen (microspore mother cell).

The formation of microspores from a pollen mother cell through meiosisis called
microsporogenesis.

As the anthers mature and dehydrate, the microspores dissociate from each other and
develop into pollen grains.

Each microsporangium contains thousands of pollen grains .They are released with the
dehiscence of anther

Pollen grain (male gametophyte)

Generally spherical generally 25-50, μm in diameter.
Cytoplasm is surrounded by a plasma membrane. A pollen
grain cell wall is made up of exine and intine.

Exine: The hard outer layer. Made up of

sporopollenin (highly resistant organic material).
It can withstand high temperature and strong
acids and alkali. [Karnataka NEET 2013]
Enzymes cannot degrade sporopollenin. Intine: The inner wall. It is a
Exine has apertures called germ pores where thin and continuous layer
sporopollenin is absent made up of cellulose and
Pollen grains are preserved as fossils due to pectin.
the presence of sporopollenin. Exine exhibits
patterns and designs.

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 Matured pollen grain contains 2 cells:

1. Vegetative Cell: It is bigger, has abundant food

reserve and a large irregularly shaped nucleus.

2. Generative cell: It is small and floats in the cytoplasm

of the vegetative cell. It is spindle shaped with dense
cytoplasm and a nucleus. In over of angiosperms,
pollen grains are shed at the 2 - celled stage In
others, the generative cell divides mitotically to give
rise to the two male gametes before pollen grains
are shed (3 - celled stage).

The shed pollen grains have to land on the stigma

before they lose viability. The viability period of pollen
grains depends on prevailing temperature and
humidity.

Viability of pollen grains of some cereals (rice, wheat

etc.) is 30 minutes within their release. Some
members of Leguminoseae, Rosaceae & Solanaceae
have viability for months.

Economic importance of pollen grains:

These are rich in nutrients. Pollen tablets are used
as food supplements. [AIPMT 2014] Pollen tablets
& syrups increase performance of athletes and
race horses.

Pollen grains can be stored for years in liquid

O
nitrogen (-196 C). They are used as pollen banks,
similar to seed banks, in crop breeding
programmes. [NEET 2018]

Pollen grains of some plants (e.g. Parthenium or

carrot grass) are allergic for some people. It leads
to chronic respiratory disorders asthma, bronchitis,
etc.

gynoecium (pistil)
It represents the female reproductive part of the flower.

It may consist of a single pistil (monocarpellary) or more than

one pistil (multicarpellary).

In (multicarpellary) the pistils may be fused together

(syncarpous) or free (apocarpous).

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 A Hibiscus
pistil.
B Multicarpellary,
syncarpous pistil
of Papaver .
c
Multicarpellary,
apocarpous
gynoecium of
Michelia.

Each pistil has three parts:

Stigma: It is a landing platform for pollen grains.

Style: It is an elongated slender part beneath the stigma.

Ovary: It is the basal bulged part of the pistil. Inside the ovary is the ovarian cavity
(locule) in which the placenta is located. Arising from the placenta are the ovules
(megasporangia). The number of ovules in an ovary may be one (wheat, paddy, mango
etc.) to many (papaya, water melon, orchids etc.)

Megasporangium (OVULE)

It is a small structure attached to the placenta by means of a stalk (funicle). The

junction where the body of ovule and funicle fuse is called hilum.

Each ovule has one or two protective envelopes called integuments. Integuments
encircle the ovule except at the tip where a small opening (micropyte) is present.

Opposite the micropylar end is the chalaza (basal part).

Enclosed within the integuments, there is a mass of cells called nucellus. Its cells
contain reserve food materials.

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 The embryo sac (female gametophyte) is found in the nucellus. An ovule
generally has a single embryo sac formed from a megaspore through meiosis.

Megasporogenesis;

monosfoiic
It is the formation of megaspores from the megaspore mother cell (MMC).

Ovules generally differentiate a single megaspore mother cell in the micropylar

region of the nucellus. It is a large cell containing dense cytoplasm and a prominent
nucleus.

The MMC undergoes meiotic division. It results in the production of 4 megaspores.

[NEET-II 2016]

Female gametophyte (embryo sac):

In a majority of flowering plants, one of the megasporesis
functional while the other three degenerate.

The functional megaspore develops into the female

gametophyte. This method of embryo sac formation from a
single megaspore is termed monosporic development.

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 The nucleus of the functional megaspore divides mitotically to
form two nuclei. They move to the opposite poles, forming
2-nucleate embryo sac.

The nuclei again divide two times forming 4-nucleate and

later the 8-nucleate stages of the embryo sac.

These divisions are strictly free nuclear, i.e. nuclear divisions are not followed
immediately by cell wall formation.

After the 8-nucleate stage, cell walls are laid down leading to the organisation of
the typical female gametophyte or embryo sac.

The 6 of the 8 nuclei are surrounded by cell walls and organised into cells.
Remaining 2 nuclei (polar nuclei) are situated below the egg apparatus in the large
central cell.
distribution of the cells within the embryo sac:

A typical mature embryo sac is 8-nucleate and 7-celled.

3 cells are grouped at the micropylar end form egg apparatus.

It consists of 2 synergids and one egg cell.

Synergids have special cellular thickenings at the micropylar

tip called fillform apparatus. It helps to guide the pollen tubes
into the synergid.

3 cells at the chalazal end are called the antipodals.

The large central cell has two polar nuclei.

pollination
It is the transfer of pollen grains from the anther to the
stigma of a pistil.

Some external agents help the plants for pollination.

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depending upon the source of pollen, pollination is 3 types:

a) autogamy (self - pollination):

In this, pollen grains transfer from the anther to stigma of the same flower.
In flowers with exposed anthers & stigma, complete autogamy is rare.
Autogamy in such flowers requires synchrony in pollen release and stigma
receptivity. Also, anthers & stigma should lie close to each other. Plants like
Viola (common pansy), Oxalis & Commelina produce 2 types of flowers:

1 2
Cleistogamous Flowers: They do not
Chasmogamous Flowers: open at all. Anthers & stigma lie
They are similar to flowers close to each other. They are
of other species with autogamous. [Karnataka NEET 2013]
exposed anthers and stigma. When anthers dehisce in the flower
pollen grains buds, come in contact
with stigma for pollination.
Cleistogamous flowers produce
assured seed-set even in the
absence of pollinators. [NEET 2013]

b) geitonogamy:

In this, pollen grains transfer from the anther to the stigma of

another flower of the same plant. [AIPMT 2014] It is functionally
cross-pollination involving a pollinating agent. But it is genetically
similar to autogamy since the pollen grains come from the same
plant.

c) xenogamy:
crass
pollination
In this, pollen grains transfer from anther to the stigma of a different
plant. It brings genetically different pollen grains to the stigma.

Agents of pollination
1. Abiotic agents (wind & water)
pollinated by wind (anemophily)
More common abiotic agent.

Wind pollinated flowers often have a single ovule in each ovary and numerous
flowers packed into an inflorescence. [AIPMT 2012]

Eg: Corn cob – the tassels are the stigma and style which wave in the wind to
trap pollen grains. Wind-pollination is quite common in grasses. [AIPMT 2011]

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 ways for effective pollination:
The flowers produce enormous amount
of pollen.

The pollen grains are light and non-

sticky so that they can be transported
in wind currents.

They often possess well-exposed

stamens (for easy dispersion of pollens
into wind currents).

Large, feathery stigma to trap air-

borne pollen grains.

pollinated by water (hydrophily):

It is quite rare. It is limited to about 30

genera, mostly monocotyledons. E.g.
Vallisneria & Hydrilla (fresh water),
Zostera (marine sea-grasses), etc.

As against this, water is a regular mode of transport for the male gametes among
the lower plants. It is believed, particularly for some bryophytes & pteridophytes, that
their distribution is limited because of the need for water for the transport of male
gametes and fertilisation.

In Vallisneria , the female flower reaches the surface of water by the long stalk and
the male flowers or pollen grains are released on to the surface of water. They are
carried by water currents and reach the female flowers. [Odisha NEET 2019]

In sea grasses, female flowers remain submerged in water. Pollen grains are long and
ribbon like. They are carried inside the water and reach the stigma.

The pollen grains of most of the water-pollinated species have a mucilaginous

covering to protect from wetting.

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 Not all aquatic plants use hydrophily. In most of aquatic plants (water hyacinth,
water lily, etc.), the flowers emerge above the level of water for entomophily or
anemophily. [NEET 2020]

Wind and water pollinated flowers are not very colourful and do not produce nectar.

2. biotic agents (animals)

Majority of flowering plants use animals as pollinating agents. E.g. Bees,

butterflies, flies, beetles, wasps, ants, moths, birds (sunbirds & humming birds)
bats, primates (lemurs), arboreal (tree-dwelling) rodents, reptiles (gecko lizard &
garden lizard), etc.

Pollination by insects (entomophily), particularly bees is more common.

Often flowers of animal pollinated plants are specifically adapted for a

particular species of animal.

features of insect - pollinated flowers:

Large, colourful, fragrant and rich in nectar. Nectar & pollen grains are the
floral rewards for pollination.

When the flowers are small, they form inflorescence to make them visible.

The flowers pollinated by flies and beetles secrete foul odours to attract these
animals.

The pollen grains are generally sticky. When the animal comes in contact
with the anthers and the stigma, its body gets pollen grains. When it comes in
contact with the stigma, it results in pollination.

Some plants provide safe places as floral reward to lay eggs. [NEET 2017]

Eg: Amorphophallus (tallest flower of 6 feet). A moth species and the plant
Yucca cannot complete their life cycles without each other. The moth deposits
its eggs in the locule of ovary. The flower gets pollinated by moth. The larvae
come out of the eggs as seeds start developing.

Many insects consume pollen or nectar without bringing about pollination. They
are called pollen/nector robbers.

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outbreedind devices
Hermaphrodite flowers can undergo self-pollination. Continued self-pollination
results in inbreeding depression. To avoid self-pollination and encourage
cross-pollination, there are some devices in plants:

a) avoiding synchronization

Here, the pollen is released before the stigma becomes receptive or stigma
becomes receptive before the release of pollen. It prevents autogamy.

c) self - incompatibility
b) arrangement of anther &
stigma at different It is a genetic mechanism to prevent self-
positions pollen (from the same flower or other
flowers of the same plant) from fertilisation
This also prevents by inhibiting pollen germination or pollen
autogamy. tube growth in the pistil.

d) production of unisexual flowers:

If male & female flowers are present on the same plant (i.e.,monoecious, e.g.
castor & maize), it prevents autogamy but not geitonogamy. In dioecious plants
(e.g. papaya), male and female flowers are present on different plants (dioecy).
This prevents both autogamy and geitonogamy. [NEET 2017]

pollen - pistil interaction:

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 It is a process in which pistil recognises compatible or incompatible pollen
through the chemical components produced by them. [NEET-I 2016]

If the pollen is (right type), the pistil accepts it and promotes post-pollination
events. Pollen grain germinates on the stigma to produce a pollen tube through
one of the germ pores. The contents of pollen grain move into the
pollen tube. Pollen tube grows through the tissues of stigma and style and
reaches the ovary.

If the pollen is incompatible (wrong type), the pistil rejects pollen by preventing
pollen germination on the stigma or the pollen tube growth in the style.

A line continious dialogue between the pollen grain and the pistil results in the
pistil's ability to recognised the pollen to accept or reject it.

In some plants, pollen grains are shed at 2-celled condition (a vegetative cell & a
generative cell). In such plants, the generative cell divides and forms the two
male gametes during the growth of pollen tube in the stigma. [AIPMT 2007]

In plants which shed pollen in the 3-celled condition pollen tubes carry 2 male
gametes from the beginning.

Pollen tube reaches the ovary, then enters the ovule through micropyle and then
enters one of the synergids through the filiform apparatus. The filiform apparatus
present at the micropylar part of the synergids guides the entry of pollen tube.

A plant breeder can manipulate pollen-pistil interaction, even in incompatible

pollinations, to get desired hybrids.
artificial hybridisation:

It is a crop improvement programme in which desired pollen grains are used

for pollination.

This is achieved by following techniques:

1. Emasculation: Removal of anthers from the bisexual flower bud of female

parent before the anther dehisces using a pair of forceps.

2. Bagging: Here, emasculated flowers are covered with a suitable bag (made up
of butter paper) to prevent contamination of its stigma with unwanted pollen.
When the stigma attains receptivity, mature pollen grains collected from
anthers of the male parent are dusted on the stigma. Then the flowers are
rebagged and allowed to develop the fruits.

For unisexual flowers, there is no need for emasculation. Female flower buds
are bagged before the flowers open. When the stigma becomes receptive,
pollination is carried out using the desired pollen and the flower rebagged.

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 double fertilisation
After entering one of the synergids, the pollen tube releases the 2 male gametes
into the cytoplasm of the synergid. One male gamete moves towards the egg cell
and fuses with its nucleus. This forms the zygote (a diploid cell).
syngamy
The other male gamete moves towards the two polar nuclei located in the
central cell and fuses with them to produce a triploid primary endosperm
nucleus (PEN). As it involves fusion of 3 haploid nuclei it is called triple fusion.

Since 2 types of fusions (syngamy & triple fusion) take place in an embryo sac, it
is called double fertilisation. As, it is a events unique to flowering plants. [NEET
2019]

The central cell after triple fusion becomes the primary

endosperm cell (PEC) and develops into the endosperm while the
zygote develops into an embryo.

post - fertilisation; structures & Events

Post - Fertilisation Events: Endosperm & embryo development,
maturation of ovule(s) into seed(s) & ovary into fruit.

endosperm development

The primary endosperm cell divides repeatedly and forms a triploid endosperm
tissue.

Endosperm cells are filled with reserve food materials. They are used for of the
developing embryo.

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 In the most common type of embryo endosperm development, the
PEN undergoes successive nuclear divisions to give rise to free nuclei.
This stage is called free - nuclear endosperm. The number of free
nuclei varies greatly.

The formation of cell walls causes the endosperm to become cellular.

In coconut, the surrounding white kernel is the cellular endosperm,
while the soft coconut water is a free-nuclear endosperm (made up of
more than thousands nuclei). [NEET-I 2016]

embryo development
Embryo develops at the micropylar end of the embryo
sac where the zygote is situated.

Most zygotes divide only after the formation of

certain amount of endosperm. This is an adaptation to
provide nutrition to the developing embryo.

Though the seeds differ greatly, the embryogeny

(early developments) is similar in monocots & dicots.

The zygote gives rise to the proembryo and

subsequently to the globular heart - shaped and
mature embryo.

dicotyledonous embryo

It has an embryonal axis and 2 cotyledons.

The portion of embryonal axis above the level of cotyledons is

the epicotyl, which terminates with the plumule (stem tip).

The cylindrical portion below the level of cotyledons is

hypocotyl that terminates with the radicle (root tip). The root
tip is covered with a root cap.

monocotyledonous embryo:

They possess only one cotyledon.

In the grass family, the cotyledon is called scutellum

It is situated lateral to the embryonal axis. At its lower end, the

embryonal axis has the radicle and root cap enclosed in coleorrhiza
(an undifferentiated sheath).

Portion of embryonal axis above the level of attachment of scutellum

is the epicotyl. It has a shoot apex and a few leaf primordia enclosed in
coleoptile (a hollow foliar structure).

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 seed from ovule

Seed is the fertilised ovule formed inside fruits. It is the final product of sexual
reproduction.

It consists of seed coat(s), cotyledon(s) & an embryo axis.

The cotyledons are simple, generally thick and swollen due to storage
food (as in legumes).

Mature seeds are 2 types:

Non - albuminous seeds: Albuminous seeds: retain a part of

have no residual endosperm endosperm as it is not completely
as it is completely consumed used up during embryo
during embryo development development (e.g., wheat, maize,
(e.g., pea, groundnut, beans). barley, castor, coconut, sunflower).

Oil reserve of groundnut is present in cotyledons.

Occasionally, in some seeds(black pepper, beet etc.) remnants of nucellus are also
persistent. It is called perisperm. [NEET 2019]

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 Integuments of ovules harden as tough protective seed coats. It has a small pore
(micropyle) through which O 2 & water enter into the seed during germination.

As the seed matures, its water content is reduced and seeds become dry (10-15 %
moisture by mass). The general metabolic activity of the embryo slows down. The
embryo may enter a state of inactivity (dormancy). If favourable conditions are
available (adequate moisture, oxygen and suitable temperature), they germinate.

seed from ovary

The ovary develops into a fruit. Transformation of ovules into seeds and ovary
into fruit proceeds simultaneously.

The wall of ovary develops into pericarp (wall of fruit).

The fruits may be (e.g. guava, orange, mango, etc.) or dry (e.g. groundnut,
mustard etc.).

Fruits are 2 types:

True fruits: In most plants,

the fruit develops only from False fruits: In this, the
the ovary and other floral thalamus also contributes
parts degenerate and fall to fruit formation.
off. They called true fruits. Eg: Apple, strawberry,
Eg: Watermelon, lemon, cashew, etc.
cherry, etc.

In some species fruits develop without fertilisation. Such fruits are called
parthenocarpic fruits. E.g. Banana. [AIPMT 2015]

Parthenocarpy can be induced through the application of growth hormones. Such

fruits are seedless.

advantages of seeds:

Since pollination and fertilisation are independent of water, seed formation is

more dependable.

Seeds have better adaptive strategies for dispersal to new habitats and help
the species to colonise in other areas.

They have food reserves. So, young seedlings are nourished until they are
capable of photosynthesis.

The hard seed coat protects the young embryo.

Being products of sexual reproduction, they generate new genetic

combinations leading to variations.

Dehydration and dormancy of mature seeds are crucial for storage of seeds. It
can be used as food throughout the year and also to raise crop in the next
season.

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viability of seeds after dispersal:

In a few species the seeds lose viability within a few months. Seeds of many
species live for several years.

Some seeds can remain alive for hundreds of years. The oldest is that of a
lupine (Lupinus arcticus) excavated from Arctic Tundra. The seed germinated
and flowered after an estimated record of 10,000 years of dormancy.

2000 years old viable seed is of the date palm ( Phoenix dactylifera )
discovered during the archeological excavation at King Herod’s palace near
the Dead Sea.

In some plants fruits contain very large number of seeds. Orchid, Orobanche
and Striga fruits are one such category & each fruits contain thousands of
tiny seeds.

APOMIXIS AND POLYEMBRYONY

Apomixis is the production of seeds without fertilisation. E.g: Some species of
Asteraceae and grasses. [NEET 2016]

It is a form of asexual reproduction that mimics sexual reproduction.

development of apomictic seeds:

In some species, the diploid egg cell is formed without reduction division and
develops into the embryo without fertilisation.

In many species (e.g. manyCitrus & Mango varieties) some of the nucellar cells
surrounding the embryo sac divide, protrude into the embryo sac and develop into
the embryos. In such species each ovule contains many embryos. [AIPMT 1995]

Occurrence of more than one embryo in a seed is called polyembroyony.

importance of apomixis in hybrid seed industry

If the seeds collected from hybrids are sown, the plants in the progeny will
segregate and lose hybrid characters.

Production of hybrid seeds is costly. Hence, the cost of hybrid seeds is also
expensive for the farmers.

If the hybrids are made into apomicts, there is no segregation of characters in

the hybrid progeny. Then the farmers can keep on using the hybrid seeds to
raise new crop year after year.

Additional Points

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