Sexual Reproduction in Flowering Plants

Flowers are objects of aesthetic, ornamental, social, religious and cultural

value. Flowers are morphological and embryological marvels and the sites
of sexual reproduction in angiosperms.

A typical angiospermic flower consists of four whorls of floral appendages

attached on the receptacle.

1. Calyx (consists of sepals): Non-essential whorl (sterile)

2. Corolla (consists of petals): Non-essential whorl (sterile)
3. Androecium (consists of stamens): male unit: Essential whorl
(fertile)
4. Gynoecium (consists of carpels): female unit: Essential whorl
(fertile)

Pre- Fertilization – Structure and Events

Much before the actual flower is seen on a plant, the decision that the plant is going to flower has taken place. A
number of hormonal and structural transformations occur prior to
initiation of flowering. Shoot apical meristem is transformed into
reproductive meristem. Reproductive meristem grows to form
inflorescence axis over which floral primordial develops. The
primordial grow into floral buds and then flowers. In the flower, the
androecium and gynoecium differentiate and develop.

The male conceptive structure is Androecium. It comprises of whorl of

stamens.

Stamen: It has slim stalk known as Filament. At its tip is contained a

bilobed structure called anther. Usually these are bilobed anthers
containing two theca (dithecous) isolated by longitudinal groove. The
transverse section of any anther shows four sided micro sporangia
two in every flap. Microsporangia further form the pollen sac.
Contained in the pollen sac are the pollen grains.
Fig.: (a) A typical stamen;
(b) Three dimensional cut section of an anther

Development of anther:

A young anther constitutes homogenous mass of meristematic cells surrounded by epidermis. The pollen sacs
develop hypodermally at the four corners of the anther from a strip of archesporial cells (archesporium). These
cells divide periclinally to form primay parietal cells (PPC) on outer side and primary sporogenous cells (PSC) on
inner side. Then PPC (outer side) further divides anticlinally and periclinally to form 3-5 layers of anther wall while
inner PSC form pollen mother cells or microspore mother cells (2n).

Structure of Anther
The fertile portion of stamens is called anther. Each anther is usually made up of two lobes connected by a
connective. In turn each anther lobe contains two pollen chambers placed longitudinally. All pollen grains of an
anther lobe remain united in a sac called pollinia.

The pollen sacs are surrounded by following 4 layers – Epidermis, Endothecium, Middle layer and Tapetum.

A typical angiospermic anther is bilobed with each lobe having two theca. The anther is a four sided structure
consisting of four microsporangia located at the corners, two in each lobe.

In a transverse section, a typical microsporangium is circular in outline and is surrounded by four well layers the a)
Epidermis b) Endothecium c) Wall Layers d) Tapetum.

a) Epidermis : The epidermis is one celled thick, the cells present between the pollen sacs are thin walled and their
region is called as stomium which is useful for dehiscence of pollen sacs.

b) Endothecium : It is present below the epidermis and expands radially with fibrous thickening, at maturity these
cells lose water and contract and help in dehiscence of pollen sacs.

c) Wall Layer : Beneath the endothecium, there are thin walled cells, arranged in one to five layer , which also help
in dehiscence of anther.

d) Tapetum : The innermost wall layer is tapetum, the cells are large, with thin cell walls, abundant cytoplasm and
have more than one nuclei. Tapetum is a nutritive tissue which nourishes the developing pollen grains.

The centre of microsporangium consists of sporogenous tissue which undergoes meiotic division to form
microspore tetrads. This process is known as microsporogenesis.

T.S of an anther and an enlarged view of one microsporangium

Microsporengiogenesis: It is the procedure of development of microspores from dust mother cells through the
process of meiotic cell division. The microspores framed are a group of four cells called Microspore Quadruplicate.
As there is further development in the anther the moisture from the anther also dries out. The microspores
separate and form into dust grains.
Sporogenous Tissue: When the anther is young, a group of compactly arranged homogenous cells called the
sporogenous tissue occupies the centre of each microsporangium.
Pollen Grains:

Pollen Grains or Dust Grains are round in shape. It consists of two layers: external called Exine and internal called
Intine. The exine is comprised of a natural compound sporopollenin. It is hard and resistance natural
compound.No one protein is known to debase this protein. The exine consists of a typical pore called germ pore.

As the dust grain develops, it contains two cells: a greater Vegetative Cell and minute Generative Cell. The
generative cell isolates mitotically into two male gametes.

Pollen grains are microscopic structures that vary in size and shape. Some are tiny orbs, while others are egg-
shaped. Although too small to see individually, they can be seen by the naked eye in large quantities.

Viewed through a microscope, a pollen grain hardly looks real. An extremely durable body, it has a tough
outer coating. This hardy coat offers great protection from the harsh outdoor environment. This is important
because inside this tough shell lie two cells: the tube cell, which will eventually become the pollen tube, and a
generative cell, which contains the male sperm nuclei needed for fertilization.

There are three main components of a pollen grain. The inside of the grain is made up of cytoplasm. This fluid
medium houses the aforementioned living cells, keeping them moist and alive. The outer shell is made up of two
layers. The inside layer is aptly named the intine (think interior). It is composed partly of cellulose, a common
component in the cell walls of plant cells.

The tough-as-nails outer layer is known as the exine (think exterior). This highly sophisticated and complex outer
layer is rich in a compound known as sporopollenin. Waterproof, resistant to deterioration and very stiff, this shell
is basically one of nature's most advanced polymers. It ensures that the tender cells inside have a strong chance of
survival.

In addition, often times the exine has folds, creases and spikes rising from its surface. Like extra armour, these
features add to the protective nature of this layer. They also play an important role in the mobility of the grains,
making it more likely that they will stick to the legs of insects as well as catch the wind.

Pollen is produced and transported in slightly different ways depending on whether the plant is an angiosperm or
a gymnosperm. Angiosperms are flowering plants, and gymnosperms are non-flowering. Let's first take a look at
pollen in angiosperms.
In angiosperms, pollen is produced by the anther, which sits on the filament in the center of the flower. Look
closely and you will most certainly see the fine yellow powder on each anther.
In order to complete fertilization, pollen must make its way to another plant. Since pollen cannot move on its own,
it must rely on other methods of transport.

Pollen Viability:
The period for which the pollen grains retain the ability to germinate on landing on the stigma is called as pollen
viability. It is highly variable and depends on prevailing temperature and humidity.

Pollen viability periods in plants:

(i) 30 minutes – Rice, wheat (cereals)
(ii) Several months – Leguminosae, Rosaceae and Solanaceae

Pollen Banks: Storage of pollen grains for years in liquid N2(-196⁰C) for later use in plant breeding programmes,
known as cryopreservation. These centres for storage of pollens are called pollen banks.
 Transformation of microspore mother cell (2n)
into pollen grain (n) is the first gametophytic
structure. On germination pollen grain develops
into male gametophyte which starts even before
pollination

The entire process of development of male

gametophyte can be divided into two phases:
a. Pre-pollination development and

b. Post-pollination development.

a. Pre-Pollination Development:
Pollen grains show in-situ germination, i.e.
germination starts in pollen sac. This is known as
precocious germination. In this process a large
central vacuole is formed which pushes the
nucleus to one side. Now the nucleus undergoes
mitosis giving rise to two daughter nuclei. Out of
the two cells so formed, the smaller one is called
generative cell and the larger one is called
vegetative cell.

Vegetative cell has plenty of cytoplasm which has

reserve food for the development of male
gametophyte. Generative cell moves to the

middle of pollen grain. At this stage pollen grain falls on the

stigma and the rest of the development takes place on the
stigma.

b. Post-Pollination Development:
As the pollen grain falls on the stigma it absorbs nutrients from
stigma through the germ pore. Due to this absorption
vegetative cell enlarges and intine moves out through the
germ pore to form pollen tube. Nucleus of generative cell and
vegetative cell migrates to the pollen tube.

Now is the time for the generative cell to divide and give rise
to two haploid, unicellular and non-motile male gametes.
Hydrolytic enzymes are secreted by the pollen tube which
dissolves the tissues of style for deeper penetration of tube.
Size of mature male gametophyte is reduced and it gets
nutrition from the tissues of style.

Fig.: Development of male gametophyte

Gynoecium

The female regenerative part of the flower is the Gynoecium.

It comprises of pistil. Gynoecium is termed in many ways:

Monocarpellary : Gynoecium comprises of single pistil. For

Example: Pea, Bean.

Multicarpellary: Gynoecium comprises of more than one

pistil.

Syncarpous: Gynoecium with combined pistil. For Example:

Tomato, Cucumber.

Apocarpous: Gynoecium with free pistil. For Example: Lotus

Vinca.

Pistil: Each pistil has three sections - the Stigma, Style and Ovary.

• Stigma: At the tip of pistil is the stigma that provides a landing stage for dust grains.

• Style: The style is the lengthened slender part underneath the stigma.

• Ovary: The basal lump part of the pistil is ovary.

The hole inside the ovary is called Ovarian Hole (Locule). The Megasporangia usually called Ovule emerge from
the placenta. The quantity of ovules inside an ovary might be one (mango, paddy, wheat) or numerous (orchids,
watermelon, papaya).

Megasporangium (Ovule): The ovule is a minute structure connected to the placenta with a stalk known as
Funicle. The ovule's body wires with funicle in the area called Hilum.

Each ovule has maybe a couple of defensive envelopes called integuments. Integuments encompass the ovule
aside from at the tip. The minute opening at the tip is called Micropyle. Chalaza is a structure present opposite to
the micropylar end. It denotes the basal part of the ovule.

The integument encloses nucellus which is a mass of cells. These cells have abundant nourishment materials.
Inside the nucleus is the embryo sac or female gametophyte. An ovule by and large has a single embryo sac
shaped from a megaspore through reduction division.
A typical anatropous ovule:

Structure of ovule (Megasporangium):

Ovule is considered to be
an integumentedmegasporangium. The ovule consists of the
stalk and the body. The stalk is called funicle. One end of the
funicle is attached to placenta and the other end to the body
of the ovule. The point of attachment of funicle with the
body is called hilum. Sometimes funicle gets fused with the
body of the ovule one side and forms a ridge known
as raphe. The body of the ovule shows two ends: the basal
end, often called the chalazal end and the upper end is
called micropylar end. The main body of the ovule is covered
with one or two envelopes called integuments. These leave
an opening at the top of the ovule called micropyle. The
integuments enclose a large parenchymatous tissue known
as nucellus. The residual part of nucellus in the mature seed is called perisperm. In the centre of the nucellus is
situated a female gametophyte known as embryo sac.

The structure of ovule can be studied under the following headings:

(a) Funicle: The stalk of the ovule by which it remains attached to placenta.
(b) Hilum: It is the junction between ovule and funicle or the point of attachment of funiculus to the body of
the ovule.
(c) Integument:The one or more protective envelops surrounding the body of ovule.
(d) Micropyle: The pore or passage present at the tip of ovule where the integument is absent.
(e) Chalaza: Opposite to micropylar end representing the basal part of ovule.
(f) Nucellus: The parenchymatous mass of tissue enclosed within the integuments and forms the body of
ovule. Depending upon the development of nucellus, ovules are of two types:
i) Crassinucellate ovule: The nucellus is well developed, eg., polypetalae
ii) Tenuinucellae ovule: The nucellus is poorly developed, eg., gamopetalae
(g) Embryo sac: It is also called the female gametophyte and is located in the nucellus. An ovule generally has
a single embryo sac formed from a megaspore.

Following are the conditions seen in ovule in relation to integuments-

(i) Unitegmic: Ovule with a single integument, e.g., sympetalous or gamopetalousdicotyledons (Compositeae) and
gymnosperms.

(ii) Bitegmic: Ovule with two integuments as in polypetalous (Archichlamydeae) dicotyledons (Papilionaceae) and
monocotyledons.

(iii) Ovule may be surrounded by three integuments (e.g., Asphodelus)

(iv) Ategmic : In some parasitic plants like Loranthus, Viscum, Santalum etc., there is no integument. Such an ovule
is called ategmic.

(v) Aril : This is a collar-like outgrowth from the base of the ovule and forms third integument. Aril is found in
litchi, nutmeg, etc.

(vi) Caruncle: It is formed as an outgrowth of the outer integument in the micropylar region.

Caruncle is common in the ovules of Euphorbiaceae. e.g., Castor (Ricinus).

Types of ovule

There are six types of ovules:

1. Orthotropous or atropous ovule(ortho-straight, tropous - turn)

The body of the ovule is erect or straight. The hilum, chalaza and the micropyle lie in a straight line e.g.Polygonum.

2. Anatropous ovule(ana - backward or up, tropous - turn)

The body of the ovule becomes completely inverted during the development so that the micropyle lies very close
to the hilum, eg.,Gamopetalae members.

3. Hemi-anatropous or hemitropous ovule

The body of the ovule is placed transversely at right angles to the funicle. The micropyle and chalaza lie in one
straight line e.g.Ranunculus.

4. Campylotropous
ovule(kampylos - curved)

The body of the ovule is curved or

bent round so that the micropyle
and chalaza do not lie in the same
straight line. e.g. Leguminosae.

5. Amphitropous ovule

The curvature of the ovule is very

much pronounced and the
embryosac also becomes curved
e.g.Allismaceae, and Butomaceae.

6. Circinotropous ovule

The nucellus and the axis are in the

same line in the beginning but due
to rapid growth on one side, the
ovule becomes anatropous. The
curvature continues further and
the micropyle again points upwards (e.g.) Opuntia.
Megasporogenesis

The procedure of formation of megaspores from the megaspore mother cell is known as megasporogenesis.
Ovules for the most part separate a single Megaspore Mother Cell (MMC) in the micropylar area of the nucellus.
The MMC experiences meiotic division.

The dominant part of flowering plants, one of the megaspores is utilitarian while the other three deteriorate. This
functional megaspore forms into the female gametophyte.

(Note - The nucellus is by nature diploid. MMC is diploid; the utilitarian haploid, female gametophyte is haploid.)

The core of the utilitarian megaspore divides mitotically to shape two nuclei. These two move to the inverse
shafts, forming the 2-nucleate embryo sac. Two more successive mitotic atomic divisions result in the
arrangement of 8-nucleate phases of the embryo sac.

One nucleus from every shaft comes to centre of embryo sac. They intertwine to frame secondary nucleus. The
three nuclei at the chalazal end shape the antipodal.

The three nuclei at the micropylar end sort out in to egg apparatus. They are recognized in to two Synergids with
an egg between them. The angiosperm embryo sac, at development, however 8 nucleate but are 7-celled.

Female Gametophyte or Embryo sac:

P.Maheshwari classified the embryo sac on the basis of number of megaspore nuclei participating in embryo sac
formation into following types:

Monosporic embryo sac: The formation of embryo sac from single megaspore is termed monosporic
development. Eg., Polygonum, Oenothera

Bisporic embryo sac: Two megaspore nuclei tale part in development of embryo sac. Eg., Allium, Endymion.

Tetrasporic embryo sac: All the four megaspore nuclei take part in development of embryo sac. Eg. Adoxa,
Plumbago, Drusa, Fritillaria, Paenaea, Plumbagella, Peperomia.
Based on the number of megaspores, embryo sacs can be divided into three types: monosporic, bisporic, and
tetrasporic. In the monosporic, or Polygonum-type embryo sac, meiosis of the diploid megaspore mother cell in
the nucellus produces four haploid megaspores. Three of the megaspores, usually those at the micropylar end of
the nucellus, subsequently undergo programmed cell death, leaving only one functional megaspore.
In bisporic embryo sacs, meiosis produces only two megaspores, each containing two haploid nuclei, due to the
absence of cytokinesis and cell plate formation following the second meiotic division. The megaspore nearest the
micropyle then undergoes programmed cell death, leaving a single functional megaspore with two haploid nuclei.
In tetrasporic embryo sacs, cell plates fail to form after both meiotic divisions, resulting in a single four-nucleate
megaspore. All three patterns give rise to a single functional megaspore that contains either one (monosporic),
two (bisporic), or four (tetrasporic) haploid nuclei. Note that the nuclei of the bisporic and tetrasporic embryo sacs
are not genetically identical as they are in monosporic embryo sacs, because they arise from two or four different
meiotic products.

Development of monosporic embryo sac -

In majority of angiosperms, one of the megaspore is functional while the other three degenerate. Only the
functional megaspore (n) develops into the female gametophyte. This process of embryo sac formation from a
single megaspore is termed monosporic development, also known as Polygonum type development which is
found in 80% of flowering plants.

The nucleus of chalazal functional megaspore (4th from micropyle) divides mitotically to form two nuclei
which move to opposite poles, forming the 2-nucleated embryo sac. Two more sequential mitotic nuclear divisions
result in the formation of the 4 nucleate and later the 8-nucleate stages of the embryo sac. One nucleus from each
pole moves to the middle and they form polar nuclei. These mitotic divisions are strictly free nuclear, i.e., nuclear
divisions are not followed immediately by cell wall formation. At this stage, following changes occur:
a) Three of the nuclei (n) get organised as cells at micropylar end forming egg apparatus. One is the egg cell
(n) and two are synergids (n).
b) Three nuclei get organised as antipodal cells (n) at chalazal end.
c) Two nuclei in the centre are called polar nuclei (n).
This constitutes a 7-celled and 8-nucledated embryo sac.

The female gametophyte (embryo sac) develops from a single functional megaspore. This megaspore
undergoes three successive mitotic divisions to form eight nucleate embryo sacs.

The first mitotic division in the megaspore forms two nuclei. One nucleus moves towards the micropylar end
while the other nucleus moves towards the chalazal end. Then, these nuclei divide at their respective ends
and re-divide to form eight nucleate stages. As a result, there are four nuclei each at both the ends i.e., at the
micropylar and the chalazal end in the embryo sac. At the micropylar end, out of the four nuclei only three
differentiate into two synergids and one egg cell. Together they are known as the egg apparatus. Similarly, at
the chalazal end, three out of four nuclei differentiates as antipodal cells. The remaining two cells (of the
micropylar and the chalazal end) move towards the centre and are known as the polar nuclei, which are
situated in a large central cell. Hence, at maturity, the female gametophyte appears as a 7-celled structure,
though it has 8 nucleate.

Organisation of Embryo sac:

a) Synergids or helper cells

or cooperative cells: These cells
generally possess a micropylar
nucleus and a chalazalvacuole.
The lectron microscopic studies
have revealed that the synergids
lack a cell wall on their chalazal
side at maturity. They are
characterized by the presence of
a filiform apparatus at the micropylar tip. It is in the form of finger like projections, each projection
comprising a core of microfibrils enclosed in a sheath. Usually one synergid starts to degenerate just with
pollination. The synergids perhaps secrete some chemotropic substance and thus, direct the pollen tube
growth inside the embryo sac.
b) Egg: The egg shows cytoplasmic polarity opposite to synergid and its wall is thicker at the micropylar end.
Usually the gg has a micropylar vacuole and a chalazal nucleus. Plasmodesmata connection is present in
between egg and synergids.
c) Antipodals or vegetative cells: These are vegetative cells of embryo sac. In most of the plants there are
three antipodal cells.
d) Central cell: it is the largest cell of the embryo sac. It initially contains two polar nuclei which fuse just
before fertilization to form a secondary nucleus or definitive nucleus (2n).
Pollination: Transfer of dust grains to the stigma of a pistil is called Pollination.

Type of Pollination:

• Autogamy: Transfer of dust grains from the anther to the stigma of a similar flower is called autogamy. In such
flower anthers and the stigma lie near each other with the goal that self-pollination can happen.

Chasmogamous flowers and Cleistogamous flowers: Some plants produce two sorts of flowers. For
Example: Viola (Regular Pansy), Oxalis, and Commelina. Flowers with uncovered anthers and stigma are
known to be chasmogamous flower. Flowers which don't open are known as Cleistogamous Flowers.

• Geitonogamy: Transfer of dust grains from the anther to the stigma of the diverse flower but of the similar plant
is called Geotonogamy. Hereditarily it is like autogamy on the grounds that the dust grains originate from the
same plant.

• Xenogamy: Transfer of dust grains from anther to the stigma of an alternate plant but of the same species.

Agents of Pollination: Plants utilize two abiotic (Wind and Water) and one biotic (Animals) operators to
accomplish pollination. Majority of plants utilize biotic operators for pollination. Just a little extent of plants
utilizes abiotic operators.

• Anemophily: The pollination happened by the impact of wind as an operator is called Anemophily.

• Hydrophily: The pollination happened by the impact of water as an operator in hydrophytes is called Hydrophily.

• Zoophily: The pollination happened by the impact of creatures as an operator is called Zoophily.

Anemophily: It is the name given to the pollination by wind. Little flowers are pollinated by anemophily.
These plants produce huge measure of dust grains. For Example: Sugar Stick, Grasses, Bamboo, Rice, and so on.
The anemophilous flowers are small and never shaded in bright color. They are unscented and never create
nectar. Pollen grains are little, dry and light. Stigmas are bushy and spread to trap dust grains floating in air.

Hydrophily: It is the pollination by water. Not all aquatic plants experiences hydrophily. When the
pollination happens underneath the water level it is called Hypohydrogamic. It found in plants like Hydrilla,
Ceratophyllum. When the pollination happens at the water level is called Epihydrogamic. It is found in plants like
Vallisneria.

In some other aquatic plants like water hyacinth (Eichhornia), water lily flower develops out of water and
pollination happens by wind or insects. In vallisneria, the sub-consolidated aquatic plant, the male flowers opens
at the surface of water. The discharged dust grains coasts on water flow. The pedicel of female flower stretches till
it achieves the water surface to contact with dust grains.

Zoophily: It is the pollination by creatures like bats, birds, insects, wasps, honey bees and so on. Majority
of plants are pollinated by insects.Theentamophilous flowers have some trademark highlights:

The flowers are fragrant and overlook scent. For Example: Jasmine, Rose.
The flowers arecolorful to attracted insects.
The nectar organ produces nectar to encourage visiting insects.
The dust grains are sticky or spiny. It attaches to the group of visiting insects.
To catch dust grains, the stigma has to be sticky.
On the basis of various sorts of creatures involved in pollination, they are grouped into:

• Entamophily: The pollination happened by the impact of insects as an operator is called Entamophily.

• Ornithophily: The pollination happened by the impact of birds as a specialist is called Ornithophily.

•Chirapterophily: The pollination happened by the impact of bats as a specialist is called Chirapterophily.

Yucca plant is pollinated by moth. The moth and yucca can't finish their existence without each other. The
moth drills an opening in ovary to lay its eggs. At that point it gathers dust grains from a few flowers and pushes in
the honor end of stigma. After fertilization and seed improvement, the hatchlings of moth eat developing seeds.
The unconsumed seeds are scattered to spread. Accordingly, moth can't sustain life without moth and yucca can't
imitate sexually without moth.

Inbreeding: The fertilization that happens within the single plant is called inbreeding. Larger part of plants is
androgynous. The autogamy (self-fertilization) and geitonogamy brings about inbreeding. The significant
disadvantage of self fertilization is, the offspring gets feeble and causes diminished wellness in population. It is
called inbreeding wretchedness. To stay away from these, plants create numerous structures that demoralize self
fertilization.

Outbreeding: The fertilization happens between two unique plants of same species is called out breeding. The
Xenogamy (cross-pollination) brings about outbreeding. The plants accomplished out-breeding by numerous
systems. The Unisexual Flowers are delivered in same plant or distinctive plant. In monoecious plants with
unisexual flowers, similar to castor, cucurbita, maize it maintains a strategic distance from autogamy. In diecious
plants like papaya, mulberry, it brings about xenogamy.

Self-Sterility or Incompatibility: In this dust of the flower has no fertilizing impact on stigma of a similar flower.
For Example: Passiflora, Potato.

Dichogamy: In this male and female sex organ develops at various times. Ex: china rose, lady's finger, jasmine,
Custard apple.

Herkogamy: The anther and stigma are put at various positions so that pollens are not able to achieve stigma of
same flower.

Artificial Hybridization

Pollinating the dust grains of one flower to the next flower of various plants of same species is known as Artificial
Hybridization. It is one of the significant approaches of harvest improvement program. In this strategy, the
required dust grains are utilized for pollination. The stigma is shielded from contamination with other dust grains.
It is accomplished by castration and bagging.

Emasculation: It is the expulsion of anthers/stamens from flower bud before it develops.

Bagging: It is the covering of undermined flower with margarine paper to avert contamination with different
pollens.