Prepared by K..

Gouthami,

Question: What is Megasporogenesis? Describe the development of various types of embryo

sacs (or) female gametophyte in angiosperms?

Ans: Development of Megaspore Mother Cell:

The ovule develops as multicellular placental outgrowth including the epidermal and a number of
hypodermal cells. With further development, this gives rise to nucellus and one or two integuments
from its basal region. In ovules, with two integuments, usually the inner one is formed first than the
outer one. The inner one is more delicate and inconspicuously developed than the outer one.
One hypodermal cell of the nucellus becomes differentiated from the other by its bigger size, dense
cytoplasm and conspicuous nucleus, called archesporial cell (Fig. 3.6A). The archesporial cell
divides transversely and forms an inner primary sporogenous cell and an outer primary parietal cell
(Fig. 3.6B).

The primary sporogenous cell functions as megaspore mother cell (Fig. 3.6C) and the primary
parietal cell undergoes repeated vertical divisions and forms layers of parietal cells (Fig. 3.6C).
Sometimes, the archesporial cell does not divide and directly functions as megaspore mother cell.

Megasporogenesis (Development of Megaspores):

The megaspore mother cell is diploid (2n), which undergoes meiosis and forms four haploid (n)
megaspores (Fig. 3.6D). The first division of megaspore mother cell is transverse, forming two
cells. Both the cells again divide transversely and thus four (4) haploid megaspores are formed.
The megaspores are then arranged in an axial row, called linear tetrad (Fig. 3.6D). Out of four
megaspores, only one which remains towards the chalazal end behaves as functional megaspore and
the other three which remain towards the micropylar end, gradually degenerate (Fig. 3.6E). The
functional megaspore forms the female gametophyte i.e., the embryo sac.

Megagametogenesis (Formation of female gametophyte i.e., Embryo sac):

Megaspore (n) is the first cell of the female gametophyte (Fig. 3.7A). The functional megaspore
becomes enlarged at the expense of tape tum and the nucellus and thus forms the female
gametophyte i.e., the embryo sac. Initially, the embryo sac is uninucleate and with further growth its
nucleus divides by three successive divisions and forms eight nuclei (Fig. 3.7B, C and D).
Out of eight nuclei, initially four remain towards the micropyle end and the other four towards the
chalazal end. One nucleus from each pole then moves towards the centre and forms a pair of polar
nuclei (Fig. 3.7E). These nuclei fuse together and form 2n nucleus, the definitive nucleus. It is also
known as polar fusion nucleus or secondary nucleus.
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 (Fig. 3.7F).

This type of embryo sac development is very common in angiosperms and is known as ordinary
type or normal type or Polygonum type. This type is also known as monosporic type, because, out
of four megaspores, only one remains functional and forms the embryo sac.

Other types of embryo sac development (Fig. 3.8):

Monosporic type:
1. Oenothera type:

In this type (like Polygonum type), usual linear tetrad of megaspores are formed, but instead of the
innermost one, the outermost megaspore (which is present towards micropyle) remains functional
and forms the embryo sac. The functional megaspore undergoes two successive divisions and forms
4 nuclei.

All the nuclei remain towards the micropyle. Out of four nuclei, three nuclei form the egg apparatus
(egg and two synergids) and the remaining one forms a single polar nucleus. Second polar nucleus
and antipodal cells are absent, e.g., Oenothera and other members of Onagraceae.
Polygonum type
 Oenothera type

Bisporic type:
2. Allium type:
The megaspore mother cell divides to form two cells, the upper one quickly degenerates. The lower
one then divides and forms two nuclei, distributed in the two poles. Later on, both the nuclei
undergo two successive divisions and form usual octant type of embryo sac, i.e., polygonum type.
Here two megaspore nuclei take part in the development of embryo sac i.e., bisporic type, e.g.,
Allium, Scilla, Trillium etc., of Liliaceae.
Tetrasporic type:
3. Peperomia type:
The megaspore mother nucleus undergoes meiotic division and forms four nuclei which remain
crosswise in the embryo sac without any wall. All the nuclei undergo two successive divisions and
form 16 nuclei which remain dispersed inside the sac. Later on, out of 16 nuclei, egg and one
synergid remain at the micropylar end, six antipodal cells towards the chalazal end, and the rest
eight at the centre forming polar nuclei, e.g., Peperomia of Piperaceae etc.

4. Penaea type:
Like Peperomia type, 16 nuclei are formed, those remain crosswise in the embryo sac. Later on, the
nuclei are distributed in a different manner. The egg and two synergids remain at the micropylar
end, three nuclei at the chalazal end, and four at the centre and three each on the two side walls,
e.g., Penaea of Penaeaceae.
5. Drusa type:
Like Peperomia type, initially four megaspores are formed, these are distributed in different ways.
One megaspore remains towards the micropyle, and the rest three at the chalazal end.

All the nuclei undergo two divisions and form 16 nuclei, out of which four nuclei remain towards
the micropyle and the rest twelve at the chalazal end. In the mature embryo sac, egg and two
synergids remain towards the micropyle, two (one from each pole) at the centre and the rest eleven
at the chalazal end, e.g., Drusa oppositifolia of Apiaceae.
6. Fritillaria type:
Like Drusa type, out of four nuclei formed, one nucleus remains towards the micropyle, and the rest
three at the chalazal end. The chalazal nuclei fused together and form 3n nucleus. Both the cells
thus undergo one mitotic division and again form a tetrasporic stage. Out of four nuclei, two remain
at each pole.
All the nuclei then undergo mitotic division and form eight nuclei. Out of four haploid nuclei at the
micropyle, one egg and two synergids are formed, those remain at the micropylal end; three triploid
nuclei at the chalazal end and one from each pole remain at the centre (one haploidand the other one
triploid), e.g., Fritillaria, Tulipa and some other members of Liliaceae.

7. Plumbagella type:
It is like Fritillaria type which forms 1st and 2nd tetrasporic stage with two haploid nuclei at the
micropyle and two triploid nuclei at the chalazal end of the embryo sac. Later on, the nuclei are
distributed in such a way that the egg is at the micropyle, one triploid nucleus at the chalazal end
and one triploid plus one haploid nuclei at the centre, e.g., Plumbagella of Plumbagellaceae.
8. Plumbago type:
It is like Penaea type where firstly four nuclei are formed followed by eight nucleated embryo sac.
The two nuclei at each side (four sides) remain crosswise. Later on, four nuclei, one from each side,
become aggregated in the centre. The nucleus at the micropylar end behaves as egg, e.g., Plumbago
of Plumbaginaceae.

9. Adoxa type:
In this type, the megaspore mother nucleus divides meiotically into four nuclei arranged two at each
end. Both the nuclei — further undergo mitotic division and thus eight nuclei are formed. Like the
normal type i.e., Polygonum type, one egg and two synergids remain at the micropylar region, three
antipodal cells at the chalazal end and two nuclei remain in the centre, e.g., Adoxa, Sambucus of
Caprifoliaceae.

Question: Describe the structure of mature embryo sac?

Ans: A tyical mature embryo sac is a 7- celled structure. It shows three components, viz., egg
apparatus, antiodals and central cell.

Egg apparatus: The egg apparatus is situated towards the microphylar end of the embryo sac. It
consists of an egg and two synergids.

Egg: It is the central cell of the egg apparatus. It is surrounded by its own wall. It encloses
nucleus towards lower side and a large vacuole towards upper side. Organelles like mitochondria,
golgi apparatus, plastids, E. R are concentrated around the nucleus.

Synergids: they are elongated cells present on either side of the egg. They are surrounded by
incomplete wall which is distinct near the micropylar end and disappears towards the chalazal end.
The xhalazal end lacks a wall, instesd shows two membranes. The cells are hooked towards the
micropylar end. The cell consists of nucleus at the upper end and a vaxuole at the lower end. The
apical region of each cell shows longitudinally oriented fibre- like structure called filiform appa-
ratus(habermann, 1906). the filiform apparatus is rich is polysaccharides and may be aiding the
synergid in absorption and transportion of materials from nucellus into embryo sac.

The synergids are ephemeral structures and degenerate shortly after the entry of pollen tube into the
embryo sac. The synergids play an important role in directing the growth of the pollen tube by
secteting chemically active substances.

Antipodal cells: The antipodal cells are 3 in number, situated towards the chalazal end of the
embryo sac. The nucleus of each cell undergoes endo-duplication to becomes polyploid. The
cytoplasm is rich in ER, mitochandria, plastids and dictyosomes. The antipodal cells are short lived
and degenerate after fertilization. The cells play a role in nutrition of the embryo sac.

Central cell: It is the largest cell of the embryo sac. It contains a large central vacuole which is
filled with sugars, aminacids and inorganic salts. It shows two polar nuclei, which are large with
prominent nuclei. The nuclei are present either at the centre of the cell or close to the egg appa-
ratus. The cytoplasm is rich in E.R, mitochondria, dictyosomes and plastids with starch grains. The
two polar nuclei fuse before fertilization to form diploid secondary nucleus.
Question: Write about Pollen- Pistil interaction?
Ans: The interaction between pollen and pistil parts( stigma, style and ovary) for ppollination and
fertilization may be positive or negative ( compataible or incompatible).
Interaction types:
a) Pollen x Stigma b) Pollen x Style c) Pollen x Ovary

a) Pollen x Stigma interaction: Stigma is the receptive part for pollen. It provides water or other
media for pollen germination. Secretions mainly contain lipid and phenolic compounds. Wet
stigmas( Petunia) secrete exudates and dry stigmas (cotton) do not secrete exudates. Dry stigmas
are covered by a hydric protein sheath called Pellicle. Sometimes, stigmas are covered by cuticle
(Brassica, Arabis).

Pollen- Stigma interaction can be known well in plants, which exhibit sexual of self-
incompatability. Green reported that pollen upon hydration( by stigma) release hydrolytic enzymes
which play key role in pollen – stigma interactions. Pollen wall and its protein contents also play
important role in this interaction. The most distinct response of the stigma to an incompatible
pollen is the development of a collase plug. Recognition of compatible pollen and rejection of
incompatible pollen by pistil are determined at molecular level through physiological and
biochemical changes. Rejection reaction occurs on stigma in SSI systems. Pollen wall proteins,
which induce incompatability are present in intine(GSI plants) and in exine(SSI systems).

b) Pollen x Style interaction: Pollen tubes growing through solid styles(petunia hybrida) or
hollow styles(Lilium) or through compatible or incompatible styles show noted differences in their
growing tips. The rejection reaction occurs in the style in GSI systems after the pollen tube has
grown to about two-thirds the length of the style. Recognition factor is contributed by the male
gametophyte. Studies in Petunia and Lycopersicum proved that incompatability proteins are related
to pollen tubes not stylar part.

Some common abnormal behaviours of the pollen tubes in an incompatible style are:
a) Retardation of pollen tube growth, b) Branching of pollen tube c) increased number of callose
pegs d) Higher activity of peroxidase.
Question: Describe the double fertilization in Angiosperms?

Ans:- The fusion of male and female gametes during the sexual reproduction in angiosperms is
called fertilization. The male gametes are produced inside the pollengrains and the female gametes
inside the embryo sac. The pollen are normally received by the stigma.
The mature anthers dehisces and liberate the pollengrains. They are carried away by different
agencies are usually 2- celled (one generative cell and another one is vegetative cell). After reaching
the stigmatic surfaces, the pollengrains begin to germinate. They absorb the secretions of the
stigma and increase in size. The germination of the pollen may occur either immediately or it take
few hours to a few days of their coming over the stigma. During germination, the intine grows out
through the germpore in the form of a long tube like structure called the pollen tube. Normally only
one tube is produced form each pollen(monosiphonous). But in some (Malvaceae, Cucurbitaceae
and Campanulaceae) more than one tube may be formed(Polysiphonous). In Althaea rosea 10 tubes
and in Malva neglecta as many as 14 tubes are reported. However, only one of them makes further
progress while others gradually disappears sometimes. The pollen tube may divide into one or
more branches(Amentiferae). In plants whose pollengrains are united into tetrads or into pollinia,
several pollen tubes are produced at sometimes. When a pollen is formed, most of the contents of
pollengrains pass intoit. The cytoplasm is mostly restricted to apocal region which is involved in
rapid growth. It is rich in organelles like amyloplasts and lipid bodies. The vegetative, generative
nucleus are also in this apical region. The wall of tube may be thick or thin and is formed of
cellulose and pectin.

After, the tube has emerged from the pollengrain it may its way through the stigmatic surface and
enters into the tissue through the style. The subsequent course of the tube depends upon nature of
the style. The style may be solid(Petunia, Gossypium) or hollow (Lilium, Monocots). When the
style is hollow pollen tube grows on the surface of the cells lining the style canal. The canal cells
are secretory in function and provide nutrition for the growing pollen tube. But when the style is
solitory the pollen tube pushes its way downward through the intercellular spaces between the cells
of stylar tissue. In some cases, the pollen tube pass through the cells of the style. As the pollen tube
grows through the intercellular spaces in the stylar region, some enzymes are secreted from its tip
bringing about the dissolution of middle lamellae and widening of the spaces between the cells.

Entry of pollen tube into ovule: After reaching the ovary, the pollen tube tries to enter in the
ovule. A pollen tube may enter into the ovule either through the microphyle, chalaza or funiculus.
Depending upon this three types are recognised.
Porogamy: In large number of plants, the ollen tube enters the ovule through the micropyle. This
condition is known as porogamy.
Chalazogamy: In some plants, the pollen tube enters the ovule through the chalaza. This condition
is known as chalazogamy. e.g. Casuarina, members of Amentiferae.

Mesogamy: In some Cucurbitceae members, the pollen tube penetrates into the ovule either passing
through the funiculus or integuments(Cucurbita). This is known as mesogamy.

Entry of pollen tube into embryo sac: irrespective of the place of entry into the ovule, the pollen
grows the nucellar tissue and enters the embryo sac only at micropylar end. Earlier works described
three modes of pollen tube entry into the embryo sac they are: i)through the synergid (or) ii)
between the egg and synergid (or) iii) between the embryo sac wall and a synergid.
Usually the pollengrains grows through the filiform apparatus and enters the cytoplasm of the
synergid. The contents of the pollen tube are discharged into the synergid through a terminal or sub
terminal pore. The pollen tube discharge include some amount of cytoplasm of vegetative nucleus
and two sperms. The pollen tube cytoplasm not mix with the cytoplasm of the synergid. The
synergid cytoplasm is towards the microphylar end and that of pollen tube towards the chalaza. In
embryo sacs, where synergid is about egg itself develops filiform apparatus and receives the egg
e.g. lumbago capensis.

The male gametes are true cells and each one surrounded by plasma membrane. Normally they are
spherical or ellipsoidal but sometimes it my be rod shaped or vermiform. One of the male gamate
approaches the egg and makes contact with the membrane of the egg. Finally the nucleus of the
male gamate fuses with that of the egg cell. This fusion of sperm nucleus with the egg nucleus is
known as the syngamy. The zygote resulting from syngamy gives rise to the embryo. The 2 nd male
gamete approaches the secondary nucleus of the central cell and fuses with it forming a triploid
primary endosperm nucleus(3n). The PEN lead give rise to the nutritive tissue, the endosperm. As
the fusion involves one sperm nucleus and two polar nuclei, it is called Triple fusion. Usually the
endosperm is triploid in nature. However, it may be dipolid(Allium) or polyploid (Fritillaria,
Lilium, Plumbago, Peperomia) etc.

Double fertilization: above matter is evident that both the male gametes are functional and are
involved in fertilization. One of the male gamete unites with the female nucleus of the egg bring
about fertilization. The fertilized egg is now called zygote and is a diploid structure. This was
called syngamy and was first described by strasburger. The other male gamete fused with the two
polar nuclei or the secondary nucleus resulting in triple fusion. The resultant nucleus is triploid and
is called primary endosperm nucleus. The phenomenon is known as double fertilization, as the
sperm nuclei are involved in two fertilization acts. It is a unique feature of angiosperms and was
first demonstrated by Nawaschin(1898).

Double fertilization is necessary for the production of viable seeds. The process of double
fertilization serves as stimulus to the polar nuclei, for the development of the nutritive tissue, the
endosperm. This is of advantage to the embryo which needs nourishment for further growth. In
gymnosperms, the formation of endosperm(female gametophyte) is a continous process and
develops in all the seeds, whether or not to the formation of embryo takes place. However, in
angiosperms endosperm will not be formed, its fertilization fails. Thus there will be no wastage of
energy spent in the formation of the endosperm.
 Diagram showing the sperm transfer into embryosac.
A---- entry of pollen tube into embryo sac.
B---- discharge of pollen tube contents.
C---- proposed method of gametic discharge.

Question: Describe the different types of endosperm found in the angiosperms?

(or)
What is endosperm and explain different types of endosperm?

Ans: Endosperm is a vegetative nutritive tissue present in the seeds of some plants. In angiosperms
it is a product of triple fusion and so it is a triploid tissue. Endosperm is formed from the primary
endosperm nucleus(PEN) of the embryo sac. Leemonnier(1887), Sargant(1900) and several other
embryologists regarded endosperm as second embryo modified to serve as a nutritive tissue for the
embryo.

Cytology: The ploidy of the endosperm depends upon the nature of secondary nucleus. Endos-
perm is formed from primary endosperm nucleus which is formed from double fertilization or triple
fusion. The fusion of secondary nucleus and a male gamete is called triple fusion. In Onagraceae
and Butomopsos, the secondary nucleus is represented by a single polar nucleus and so endosperm
is diploid. In Polygonum, secondary nucleus is 2n and so endosperm is typically triploid. In
Fritillaria, Plumbagella, Penaea and Plumbago the secondary nucleus is tetraploid and hence endos-
perm is pentaploid. In Paperomia, secondary nucleus is formed by the fusion of 8 nuclei and hence
endosperm is 9n. During develoment of endosperm polyploidisation of cells takes place due to
nuclear fusions or Endomitosis.

Types: In angiosperms there are 3 types of endosperm

1) Nuclear endosperm
2) Cellular endosperm and
3) Helobial endosperm
1) Nuclear endosperm: this is the most common type of endosperm. Here the embryo sac
enlarges, primary endosperm nucleus undergoes free- nuclear divisions. The nuclei remain
distrubuted in the peripheral layer of cytoplasm which surrounds the large central vacuole.
Cellurization takes place in a centripetal manner.
Eg: Phaseolus, Crotalaria.
In Crotalaria, the micropylar half becomes cellular and chalazal region remains free nuclear.

2) Cellular endosperm: Primary endosperm nucleus divides repeatedly but every nuclear
division is followed by wall formation. Hence the endosperm is cellular right from the
beginning.
e.g.: Datura, Petunia.

3) Helobial endosperm: This is common in plants of the tribe Helobiae. The primary endos-
perm cell divides into two cells- the micropylar and chalazal cells. In the micropylar cell, as
a rule, free nuclear divisions and cell formation, if any, start at a much later stage. In the
chalazal chamber, the nucleus either remains undivided or divides only a few times in which
the divisions are usually free nuclear.

Ruminate endosperm: Mature endosperm with any degree of irregularity and unevenness in its
surface contour is called ' Ruminate endosperm'. Ruminate starts at a late stage of endosperm
development. Rumination is caused by the activity of the seed coat or by the endosperm itself.

e.g.: Pasylara callarata, Annana, Myristica.

Functions of endosperm:
I) Being rich in stored food materials such as carbohydrates, proteins and fats, the
endosperm provides nourishment to the developing embryo.
II) As the embryo is embedded in the endosperm the latter protects the embryo.
III) It provides a better starting point for the seedling and helps it to establish
successfully until it becomes independent.
Question: Describe the development of Dicot embryo?
Ans: Development of the Embryo:
Side by side with the development of the endosperm, the zygote or oospore is developing the
embryo. Both dicotyledons and monocotyledons begin embryo development in the same way but,
there is considerable difference in later differentiation. In all Angiosperms the zygote (Fig. 422A)
divides to develop a. two-celled proembryo (Fig. 422B). In most plants the first wall between the
two cells is transverse while only in a few cases the first wall is more or less vertical (Piperad type,
Fig. 423A1).
Of the two cells the one near the micropyle is termed the basal cell while the
one pointing towards the centre of the embryo sac is called the terminal cell. The basal cell usually
forms the suspensor and may or may not contribute towards the future embryo in whose
development the terminal cell takes the main part.

On the basis of cellular configuration of the embryo at the four celled stage, and the part played by
the each of these cells in embryo development, dicotyledonous embryos have been classified into
the following five types:
Development of dicotyledonous embryo :
The classical example of dicotyledonous embryo development is the plant Capsella bursa-pastoris
of Cruciferae. The ovule is campylotropous so that the embryo sac and the later developed
endosperm as well as embryo are horseshoe-shaped.

Moreover the micropyle being pointed downwards, the embryo looks upside down.
Embryo development here is of the typical Onagrad or Crucifer type described above.

The oospore (Fig. 424A) divides by a transverse wall forming a large basal cell and a
smaller terminal cell (Fig. 424B). The basal cell now divides transversely while the
terminal cell divides by a vertical wall developing a │ -shaped 4-celled proembryo (Fig.
424B).

Next, the second basal cell quickly divides by a number of transverse walls giving rise to a
row of cells called the suspensor (Fig. 424C). The lowest (micropylar) cell of the suspensor
remains disproportionately large. As the suspensor increases in length, it pushes down the
terminal embryonal cells deeper into the embryo sac. The embryonal cell, meanwhile,
divides by three walls at right angles to one another giving rise to eight cells or octants (Fig.
424C).
This is called the embryonal mass. The lowest cell of the suspensor is called the hypo
physis. The embryonal mass, along with the hypophysis, divides further (Fig. 424 D & E).

Ultimately, the four terminal octants form the two cotyledons, the four micropylar octants
form the hypocotyl and the core of the radicle while the hypophysis forms the cortex and
the epidermis of the radicle as well as the root-cap (Fig. 424 F & G). In the final stage (Fig.
424H) we find the mature embryo with the plumule developed also out of the four terminal
octants. The suspensor gradually withers as the radicle is developed.

As the embryo is developing, the endosperm also keeps pace. Endosperm development, in
this case, is strictly nuclear. The endosperm mother nucleus rapidly gives rise to a number
of free nuclei which are arranged in the cytoplasm lining the periphery of the embryo sac
while the centre is occupied by a big vacuole (Fig. 424D).

Gradually, cell wall formation begins from the periphery (Fig. 424F). The antipodal cells,
in this, case, form a tissue which is short-living. The endosperm supplies nutrition to the
embryo mainly through the suspensor. As the embryo of Capsella increases, the
endosperm presses upon and sucks in plenty of food material from the nucellus which is
soon consumed.

In the final stage (Fig. 424H) the nucellus has been completely consumed and most of the
endosperm has also been consumed by the embryo. In the mature seed even this remnant
of endosperm disappears so that the seed becomes exalbuminous.
Question: Describe the development of Monocot Embryo?

Ans. The embryogeny in monocotyledons is explained in Najas lacerata. Transverse division of

the zygote results in a large basal cell(cb) and a small apical cell(ca). The basal cell, without
dividing even once, enlarges to form a single celled haustorium. Thus, the entire embryo is derived
from the apical cell which divides transversely into two cells, 'c' and 'd'. of these the cell 'd' once
again divides transversely. In this way a linear proembryo of four cells (c, m, c, cb)is formed. The
two vertical divisions at right angles to each other in the two distal cells(c,m) lead to the formation
of two superposed tiers(q,m) of four cells each. In the mean time cell 'ci' divides transversely to
give to 'n and n1' whereas cell 'n' divides vertically, 'n1' undergoes transverse division giving rise ot
two cells. The latter (p) undergoes another transverse division producing cells 'h and s'.

The quadrant 'q' divides by a periclinal division cutting a four – celled dermatogen surrounding the
four axial cells. The cells in the tier 'm' divide by vertical and transverse divisions and become two
tiered. At this stage the proembryo is slightly spherical. Now onwards it elangates appreciable due
to mainly transverse divisions in the tiers 'q,m and n' differentiates into plerome initials.

At this 8 celled stage of the tier 'q', three of the axial cells divide faster than the fourth one. This
disturbs the symmetry of the proembryo and its top becomes notched. The raidly growing portion
of the tier 'q' forms the single cotyledon and the slow growing tissue derived from the axial cell,
gives rise to the initials of Epicotyl. The radicle is organised from the derivatives of 'n'.
Thus, the epicotyl in monocotyledonous is truly a terminal structure, both epicotyl and cotyledon
arise from one and the same terminal tier. The apparent lateral position of the epicotyl is due to
early growth of the cotyledon, the epicotyl, after initiation, shows slow growth.
Question: Write an essay on Polyembryony?

Answer: Presence of more than one embryo in a seed is described as polyembryony.

Ppolyembryony was first reported by Leewenhoek. Polyembryony is very common in
gymnosperms and is and unusual feature in the species of Citrus, Mangifera. Polyembryony in
angiosperms may arise by
1. Cleavage of proembryo.
2. Formation of embryos by cells of the embryo sac other than the egg.
3. Development of more than one embryo sac within the same ovule.
4. Activation of some sporophytic cells of the ovule.

1. Cleavage polyembryony: This is quite common in orchids. The zygote or its derivatives
undergo cleavage and proliferation. As s result many embryonal primordia are formed. It is due to-
a) The zygote divides irregularly to form a mass of cells of which those lying towards the chalazal
end grow simultaneously and give rise to many embryos.
b) The proembryo gives out small buds or outgrowths which may themselves function as embryos.
c) The filamentous embryo becomes branched and each branch gives rise to an embryo.

2. Embryos from cells of the embryo sac other than the egg: Sometimes, the synergids and
antipodals may get fertilized besides the egg and polar nuclei.
e.g.: Aristolochia bracteata, Poa alpina.
This is possibel if more than two pollen tubes enter the embryo sac or due to polyspermy.
Occassionally synergids and antipodals may directly develop into haploid embryos even without
fertilization.
3. Presence of more than one embryo sac within the same ovule: Multiple embryo sacs in an
ovule may arise from:
a) Derivatives of the same megaspore mother cell.
b) Derivatives of two or more megaspore mother cells.
c) Nucellar cells.
Formation of twin embryo sacs within a ovule is known in Casuarina equisetifolia, citrus and poa
pratensis.
4. Activation of sporophytic cells of the ovule: The embryos arising from the maternal
sporophytic tissue(outside the embryo sac) are called Adventitive embryos.

The only maternal tissues which are known to form adventitive embryos are the nucellus.
e.g.: Opuntia dillenii and the integuments. (e.g.: Euonymus)

Causes of polyembryony : Several theories were put forward to explain the polyembryony.
In these, the polyembryony has been described to bybridisation, necrohormones, the effect of
recessive genes etc.

Significance:
I) Nucellar adventitive polyembryony is of great significance in horticulture.
II) The adventitive embryos provide uniform seedlings of the parental type.
III) Nuclear seedlings of Citrus furnish better clanes of orchard stock than cuttings.
IV) The nucellar seedlings show a restoration of the vigour lost after repeated propagation
by cuttings.
 V) The nucellar embryos are free from disease. So far nucellar polyembryony is the only
practical approach to raise virus-free clanes of polyembryonate citrus varieties in nature.

Question: Write a brief account of Apomixis and significance?

Ans: Apomixis may be defined as the substitution of the usual sexual reproduction by a form of a
sexual reproduction which does not involve meiosis and syngamy. Apomixis involves two main
categories.
a) Vegetative reproduction: The structural units employed for this purpose are called propagules.
The propagules include bulbs runners, suckers. e.g.: Agava, Lilium.
b) Agamospermy : The plants retained seed as the agent of propagation but the embryo is formed
by
some process in which normal meiosis and syngamy has been eliminated. It is of two types.
1. Gametophytic Apomixis : The embryo sac develops from diploid megaspore mother cell
without meiosis.
2. Adventive Embryony : The development of embryo directly from any diploid sporophytic
cell of the ovule like nucellus or integuments is known as adventive embryony.
e.g.: Citrus, Mango.
Significance of Apomixis:
1. Apomixis do not involve meiosis. Hence, segregation and recombination of
chromosomes do not occur.
2. Apomixis help in preserving desirable characters for indefinite periods.
3. Apomixis simplifies commercial hybrid seed production.
4. Adventive embryony well known in citrus, is being used to produce uniform root-stock
and virus-free scion material.

Question: Short notes on applications of palynology?

Answer: 1. Palynology helps in reconstruction of art vegetations.
2. On the basis of pollen morphology plant families are segregated into two main
groups.
a) Stenopalynous b) Eurypalynous
3. The knowledge of pollen morphology has been used to substitue many taxonomic
revisions.
e.g.: Segregation of Bombaceae (3- colporate) from malvaceae ( Pentaporate).

Question: Explain NPC system of classification of pollen grains?

Answer: Erdtman (1969) proposed NPC system of classification of pollen grains. NPC system is
based on the Number(N) Position(P) and Characters © of the apertures.

Number of apertures: On the basis of number of apertures, 9 types of pollengrains are

recognised. They are:
 Type of pollen grain Number of apertures
1. N0 ----- Atreme Nil
2. N1 ----- Monotreme One
3. N2 ----- Ditreme Two
4. N3 ----- Tritreme Three
5. N4 ___ Tetratreme Four
6. N5 ----- Pentatreme Five
7. N6 ----- Hexatreme Six
8. N7 ----- Polytreme Seven or more
9. N8 ----- Anomotreme Irregular

Position of apertures : On the basis of position of apertures, 7 types are recognised. They are:
Type of pollen grain Position of aperture

1. P0 Unknown position

2. P1 Catatreme One aperture found at the proximal pole.

3. P2 Anacatatreme Two apertures- one found at distal pole and

the other at proximal pole.

4. P3 Anatreme One aperture found at the distal pole.

5. P4 Monozonotreme Apertures arranged at the equatorial plane.

6. P5 Plezonotreme Apertures arranged in parallel zones at the

equatorial plane.
7. P6 Pantotreme Apertures arranged uniformly distributed
all over the surface.

Characters of apertures : On the basis of characters, 7 types are recognised. They are:

1. C0 ------ Pollen grains with unknown apertural characters.

2. C1---- Pollen grains with split apertures-single or three.
3. C2---- Pollen grains with 1,2 or 3 chotomocolpate apertures.
4. C3---- Pollen grains with one to many unbranched colpii.
5. C4---- Pollen grains with one to many unbranched apertures.
6. C5---- Pollen grains with one to many unbranched colporate apertures.
7. C6---- Pollen grains with one to many pores.
Question: Short notes on Exine sculpturing.
Ans: The surface of exine is characterised by ornamentation and sculpturing. The heads (capita)
of bacula fuse together to form tectum. Tectum is sculptured in characteristic ways. If the
tectum is in the form of a net, the sculpturing is called reticulate. If the tectum is in the form
of parallel rows, the sculpturing is called striate. If the tectum is in the form of irregular
rows, the sculputuring is calld ornate. If the exine shows radially directed, drumstick
shaped rods, the sculptuting is called pilate. Several such sculptures are formed. They are
useful in taxonomic studies.