Unit VI.

Plant Development

1. Formation of microgametophyte and

megagametophyte
2. Pollination
3. Double fertilization
4. Life cycle of a Flowering plant
5. Life cycle of a fern
6. Genes involved in plant development
Plants have two distinct stages in their life cycle: the gametophyte stage and the sporophyte stage.
The haploid gametophyte produces the male and female gametes by mitosis in distinct multicellular
structures. Fusion of the male and females gametes forms the diploid zygote, which develops into
the sporophyte. After reaching maturity, the diploid sporophyte produces spores by meiosis, which
in turn divide by mitosis to produce the haploid gametophyte. The new gametophyte produces
gametes, and the cycle continues.
SPOROGENESIS
• is the formation of spores
GAMETOGENESIS
• is a biological process by which haploid male and
female gametes are formed.
• This occurs in both plants and animals.
• In higher plants, there are two stages that are
involved- sporogenesis and gametogenesis
SPOROCYTE
• a diploid cell that divides by meiosis to produce
four haploid spores.
MICROSPOROGENESIS AND
THE
DEVELOPMENT OF MALE
GAMETES AND
GAMETOPHYTE OR POLLEN
GRAIN
MICROGAMETOGENESIS/POLLEN
DEVELOPMENT/POLLEN FORMATION
•Lead to the formation of male gametes
(sperm)
•occurs in two stages:

Microsporogenesis
Microgametogenesis
MICROGAMETOGENESIS/POLLEN
DEVELOPMENT/POLLEN FORMATION
• The androecium of the plant is made up of the stamen,
filament, and anthers. The pollen grains that carry the
male gametes are contained in the anthers. The anther is
anatomically divided into lobes and the lobes are further
divided into chambers called microsporangia.

• Pollens present in these microsporangia. The

microsporangia are also known as pollen sac. It is said
that the pollen sac develops from a parent cell known as
the archesporial cell. This archesporial cell divides into
outer and inner layers. The inner layer forms the
sporogenous tissue or cell that forms the pollen mother
cell.
 STRUCTURE OF ANDROECIUM
• An androecium is the male part of the flower which is composed of a long
filament and an anther attached to its tip.

• The number of stamens may vary according to the flower.

• The anther is a bi-lobed structure. Each lobe contain two pollen sacs.

• The pollen sacs contain pollen grains. The pollen grains are nourished in
the pollen sacs and when they mature they are liberated out of the sac.

• The pollen grains are very minute structures, which appear as a powdery
mass and can be found in different size and shapes.
 MICROSPORANGIUM

1. Structure in plant's male reproductive

organ which develops into pollen sac
2. Each anther has four microsporangia
3. Development of pollen takes place here
Structure of
Stamen (A)
Microsporangium/
Pollen sac
(B and C)
Structure of Microsporangium and Formation of Pollen
1. Surrounded by four wall layers - epidermis, endothecium, middle
layer, tapetum.
2. The outer three layers protect the pollens and also help in release
of pollens from anther.
3. The innermost layer, tapetum nourishes the pollen grains.
4. A group of compactly arranged homogenous cells called
sporogenous tissue are present at the centre of
microsporongium in an young anther.
5. Each sporogenous cell is known as pollen or microspore mother
cell
6. The microspores are arranged in the form of a tetrad
7. The microspores separate from each other and develop into a
pollen grains
 MICROSPOROGENESIS
• Development of microspores from
microsporocytes (microspore mother cell pollen
mother cell).
• Product is microspore
• This sporogenous tissue is surrounded by the
tapetum layer which provided nourishment to
the pollen or microspore mother cell present in
the sporogenous tissue. The microspore mother
cell is diploid and divides meiotically to form
haploid microspores or pollen. This describes
the process of microsporogenesis.
 BASIC TYPES OF
MICROSPOROGENESIS
• as determined by the timing of cytokinesis, which
is the formation of a plasma membrane and cell
wall that divides one cell into two (Figure 11.6A).
1. If cytokinesis occurs after meiosis I, then
microsporogenesis is successive (Figure 11.6B).
Successive microsporogenesis results in two
cells after meiosis I and four cells after meiosis
II.
2. If cytokinesis doesn't occur until after meiosis II,
then microsporogenesis is simultaneous (Figure
11.6C). Simultaneous microsporogenesis results
in cell formation only after meiosis II.
FIGURE 11.6. Microsporogenesis. A. Diagram showing two major types, simultaneous and successive. B. Successive
microsporogenesis (Lophiola aurea). Microsporocyte at anaphase II of meiosis. Note that cytokinesis, resulting in cell
wall formation, has occurred after meiosis
 MICROGAMETOGENESIS
• Development of pollen grains (male gametophytes) from
microspores.
• Product is pollen grain or male gametophyte or
microgametophyte
• The pollen grain is the haploid cell that is made up of two layers:
the outer exine layer which is derived from the tapetum and the
inner layer called intine. The exine is present all over the pollen
grain except in one small part from where the pollen tube
emerges post pollination. This small part is known as the germ
pore.

• The pollen grain divides into two halves- the small generative
nucleus and the larger vegetative nucleus. The generative cell
gives rise to two male nuclei whereas the vegetative cell gives
rise to the pollen tube. This explains the process of
microgametogenesis.
POLLEN GRAIN
 MICROGAMETOGENESIS
• technically occurs beginning with the first mitotic division
of the single microspore nucleus.
• One embryological character concerning this event is the
number of nuclei present in the pollen grain at the time of
anthesis, or flower maturation (Figure 11.7).
• Most angiosperms have pollen grains that are binucleate
(Figure 11.7), containing one tube cell/nucleus and one
generative cell/nucleus.
• The generative cell divides to form two sperm cells only
after pollen tube formation. In many angiosperm taxa,
however, the pollen at anthesis is trinucleate
• (Figure 11.7), caused by division of the generative cell
prior to pollen release.
FIGURE 11.7. Microgametogenesis. Pollen nucleus number at anthesis.
POLLEN DEVELOPMENT IN THE ANTHERS
A. Cross-section through an anther of Lilie (Lilium sp.) with on the left and the right side two loculi
each. In the loculi spore mother cells (SMCs) can be seen from which the four spores develop through
meiosis I and II. In between the loculi of each pair a thin layer of cells (arrow) is visible along which the
loculus can burst open atmaturity and release the pollen grains. In the middle the cross-sectioned
filament (Fi) to which the anther is attached is indicated. In the upper part the vascular bundle (v) of
the loculus can be distinguished.
B. Loculus. The lumen contains developing pollen. On the inner wall (w) of the loculus a layer
constitued of block-shaped single cells is present, the tapetum (t). The tapetum feeds the developing
spore and -later- pollen.
C. Tetrad stage during pollen development. After the two meiotic divisions the four daughter cells are
still interconnected and form a tetrad. They are still surrounded by the wall (arrow) of the original cell,
the microspore mothercell (MMC).
D. Mitotic division in the spore leading to the formation of a microgametophyte or pollen. Only the
metaphase is shown here. The chromosomes lay in the equatorial plane of the cell.
E. Nearly ripe pollen grain: visible are a vegetative cell with nucleus (VN), which later will form the
pollen tube, and a generative cell with its own nucleus (GN), which later will divide into two sperm cells.
F. Ripe pollen grain in which the texture of the outer cell wall, the exine, can be recognized. The grainy
dark purple structure in the middle of the pollen grain is the vegetative nucleus.
G. Diagram in 3 parts: Ripe pollen grain consisting of the vegetative cell (VC) and therein the smaller
generative cell (GC). After landing on the stigma (St) the pollen grain germinates and forms a pollen
tube. In the pollen tube the generative cell divides into two sperm cells (SC). The pollen tube grows to
the embryo sac (ES) and delivers the two sperm cells that are involved in double fertilization.
 SUMMARY OF POLLEN
DEVELOPMENT
Pollen grains develop in the anthers of the stamen. In the anthers mostly four, but sometimes only two
loculi are present.
In the loculi sporogenic tissue (from the Greek spora = seed and the Latin generare = produce) can be
found from which pollen develop. At the inner side of each loculus a layer of large, rectangular cells, the
tapetum (from the Greek tapes = carpet can be found.
The tapetum serves for the nutrition of the developing pollen, the deposition of cell wall material in the
outer part of the pollen grain and other compounds in and over the wall. First, free pollen mother cells
(PMC) are formed, which become spores by a meiotic division.
The meiosis involves two divisions, which lead to the formation of four daughter cells, the spores. Those
four cells are originally still interconnected and are called tetrads (Greek Tetra = four).
Later they come apart and the tapetum deposits the outer wall or exine.
The exine protects the spore against dessication, mechanical pressure and ultraviolet radiation.
Sometimes the exine layer is covered by sticky substances (pollenkitt, tryphine, elastoviscin and
sporopollenin viscin threads), which are also produced by the tapetum.
This adhesive material facilitates the attachment of pollen grains to insects, and in this way also zoophilic
pollination.
It also plays an important role in the adhesion of pollen grains to the female stigma and in the recognition
between pollen and pistil.
Also substances responsible for pollen allergy are often products originating from the tapetum.
 SUMMARY OF POLLEN
DEVELOPMENT
Mature pollen grain and pollen tube in flowering plants (REFER TO FIGURE BELOW)
In the final ripening phase of the spore a division occurs, giving rise to two cells. The
spore has now become a bi-cellular, male gametophyte or pollen grain.
Most striking is the large cell, the vegetative cell (VC) with many organelles and an own
nucleus, the vegetative nucleus (VN ).
The vegetative cell is surrounded by a thick, strongly sculptured cell wall. This large cell
encloses a second, much smaller, spindle-shaped cell, which is called generative cell
(GC) and which also bears an own nucleus, the generative nucleus (GN).
After maturation the pollen grains enter a phase of quiescence and dry out together
with the anthers. When pollen grains reach a mature stigma (the upper part of the pistil;
St ) of the same species following pollination by wind, animals or water, they take up
water.
As a result the rehydrated pollen grains can reactivate and start to germinate and form a
tubular extension, the pollen tube. The generative cell will divide once more to produce
two sperm cells (SC)
Note: plants exhibit a double fertilization); these sperm cells are thus the actual male
sex cells or gametes. The division of the generative cell into a sperm cell can take place
at various stages of development, in some species this division is already concluded in
the anthers, in other plants it takes place in the pollen tube, shortly after the germination
of the pollen grain.
A. Ripe pollen grain and B. pollen tube in tobacco
Only the nuclei, which have been stained with the fluorescing probe DAPI, are clearly visible:
The generative nucleus = NK as part of the generative cell,
The vegetative nucleus = VN, part of the vegetative cell
The two sperm nuclei = SN, belonging to the two sperm cells
Germinating pollen grain and pollen tube in Galanthus nivalis
A + B Pollen germination, C Growing pollen tube and D Generative cell (GC) in a pollen tube
 Megasporogenesis and the
development of female
gametes and gametophyte or
embryo sac in the ovary
 MEGAGAMETOGENESIS
• to form the female gametes, like the male
gametes
• occurs in two stages:

Megasporogenesis
Megagametogenesis

The word ‘mega-‘ is used instead of ‘micro-‘ in

females because the female gametes are said to
be larger in size as compared to the male
gametes.
 MEGASPOROGENESIS
• refers to the development of megaspores from
the megasporocyte, the cell that undergoes
meiosis.
• The ovules are present inside the ovary in
multiple lobes. A cell in the ovule differentiates
into a megaspore mother cell/megasporocyte.
The megaspore mother cell is diploid. This
megaspore mother cell undergoes meiosis to
form 4 haploid megaspores. 3 of the 4
megaspores degenerate and only one
megaspore is left in each ovule. This process is
known as megasporogenesis.
 MEGASPOROGENESIS
• Ovule develops as primordium and then mound of nucellus over placenta
• Initials of integuments develop from its base .
• They grow and come to surround the nucellus on all sides except at the tip or
micropylar region .
• In the hypodermal region of nucellus towards the micropyler end develops a primary
archesporial cell .
• It grows in size and develops a prominent nucleus .
• The archesporial cell often divides once into outer primary parietal or wall cell and
inner primary sporogenous cell .
• Primary parietal cell may divide one or more times .
• The primary sporogenous cell commonly functions directly as diploid megaspore
mother cell or megasporocyte .
• The megaspore mother cell ( MMC ) undergoes meiosis and forms a linear tetrad of 4
haploid megaspores .
• The process of meiotic formation of haploid megaspores from diploid megaspore
mother cell is called megasporogenesis .
• Commonly the chalazal megaspore remains functional while the other 3 degenerate .
 MEGAGAMETOGENESIS
• From the remaining functional megaspore, the
megagametophyte (also known as female
gametophyte or embryo sac) develops by three
mitotic divisions. The megaspore nucleus now
begins to divide mitotically to form 8 nuclei. 6 of
the 8 nuclei migrate to opposing poles (3 each)
while two nuclei remain at the center. The nuclei
that remain at the center are known as polar
nuclei. These polar nuclei fuse to form the
secondary nucleus. The megaspore matures
into an embryo sac. This entire process is called
as megagametogenesis.
 MEGAGAMETOGENESIS
• The functional megaspore is the first cell of female gametophyte .
• The cell enlarges and undergoes three free nuclear mitotic divisions .
• The first division produces two nucleate embryo sac .
• The two nuclei shift to the two ends and divide there twice forming four
nucleate and then eight nucleate structure .
• One nucleus from each side moves to the middle .
• They are called polar nuclei .
• The remaining three nuclei form cells at the two ends , 3 celled egg
apparatus at the micropylar end and three antipodal cells at the chalazal
end .
• The middle binucleate par organises itself into central cell .
• Embryo sac developed from a single megaspore is called monosporic .
• Maheshwari ( 1950 ) has also distinguished bisporic and tetrasporic
embryo sacs in which two and four megaspore precursors are involved
respectively in the formation of embryo sac .
Pollination
refers to the pre-fertilization event or process where
pollen grains from anther are transferred to the stigma
of a flower and subsequent germination of the pollen on
the stigma.
Double fertilization: In angiosperms, one sperm fertilizes the egg to form
the 2n zygote, while the other sperm fuses with two polar nuclei to form the
3n endosperm. This is called a double fertilization.
A L
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G F
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O
S C
P Y
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R L
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Life cycle of an angiosperm.
The mature sporophyte produces flowers. Flowers contain male (stamens)
and female (pistil) reproductive organs. In the pistil, the megaspore mother
cells in the ovule undergo meiosis. Only one of the four products of meiosis
survives and eventually develops into the eight celled female gametophyte,
containing the egg cell, the central cell and a few other cells (antipodals and
synergids). Within the anther, microspore mother cells undergo meiosis and
form four haploid microspores. Unlike in the female, each microspore
develops into a male gametophyte, or pollen grain, composed of only three
cells. The pollen grain contains two sperm cells. Pollen grains land on the
stigma surface of the pistil, grow a tube through the extracellular matrix of
the style and arrive at the ovary by bursting through one of the synergid
cells. The sperm cells cannot swim, but are delivered to the embryo sac by
growth of the pollen tube. One sperm cell fuses with the central cell to form
the primary endosperm cell and the other sperm cell fuses with the egg to
form the zygote. The zygote develops into the sporophyte. Growth of the
sporophyte is the result of mitotic divisions of cells in the root and shoot
meristems.
F
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C
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Starting with the "fern" as we recognize it (the sporophyte), the life cycle follows
these steps:
1. The diploid sporophyte produces haploid spores by meiosis, the same process
that produces eggs and sperm in animals and flowering plants.
2. Each spore grows into a photosynthetic prothallus (gametophyte) via mitosis.
Because mitosis maintains the number of chromosomes, each cell in the prothallus
is haploid. This plantlet is much smaller than sporophyte fern.
3. Each prothallus produces gametes via mitosis. Meiosis is not needed
because the cells are already haploid. Often, a prothallus produces both sperm
and eggs on the same plantlet. While the sporophyte consisted of fronds and
rhizomes, the gametophyte has leaflets and rhizoids. Within the gametophyte,
sperm is produced within a structure called an antheridium. The egg is produced
within a similar structure called an archegonium.
4. When water is present, sperm use their flagella to swim to an egg and fertilize it.
5. The fertilized egg remains attached to the prothallus. The egg is a diploid zygote
formed by the combination of DNA from the egg and sperm. The zygote grows via
mitosis into the diploid sporophyte, completing the life cycle.