FEMALE GAMETOPHYTE

MATURE EMBRYO SAC

• After the last nuclear division in the female gametophyte, the cleavages
are such that all the cells of embryo sac are formed within the wall of the
parent megaspore.
• 7-celled organisation of embryo sac (3-celled egg apparatus,three
antipodal cells,and a binucleate central cell) is most common among
angiosperms.
• The egg is universal and it is always haploid.Except for Plumbago and
Plumbagella types, the egg is always associated with two (rarely one,as in
Peperomia type) synergids.
• Antipodals are almost always present, except in Oenothera type.Their
number and ploidy is, however variable.
 Synergids
• Elongated cells present at the micropylar end of embryo sac.
• When two synergids are present they lie in contact with each other, and
partly embrace the egg.
• The wall around the synergids is incomplete. There is a distinct wall
around the micropylar one-third of the cell which thins toward the chalazal
end and, finally, disappears.
• Filiform apparatus (FA), is present at the micropylar end of each
synergid.FA is a mass of finger-like projections of the wall into the
cytoplasm.
• The cytolasm of the synergid is strongly polarized. The chalazal
region of the cell is occupied by one large or many small
vacoules.
• Large amount of cytoplasm and a prominent nucleus are present
in micropylar half of the cell.
• Synergids are ephemeral structures. In embryo sacs with two
synergids, one degenerates before or soon after the entry of
pollen tube into the embryo sac, whereas the other one, often
called the persistent synergid, degenerates shortly after the
embryo sac has received the pollen tube discharge.
• Occasionally,however, one of the synergids may persist for
a considerable period after fertilization. Its nucleus may
enlarge and show polyploidy.
• In Allium angulosum the synergid nucleus becomes
octaploid.
• In Cotula australis one of the synergids becomes swollen
and haustorial in nature.
• Quinchamalium chilense shows the most extensive synergid
haustoria.their tips elongate and breaking through the
embryo sac,grow into the stylar tissue. They reach up to
one-third the length of the style.
 Functions of synergids –
1. As the filiform apparatus (FA) increses the surface area of the
plasma membrane at the micropylar end of the synergids and
the cytoplasm near FA is rich in secretary organelles,it has
been suggested that the FA facilitates transport of substances
into and out of the synergid.
• Lot of evidence have been published to support the idea that FA
plays a role in directing pollen tube growth to the synergid cell.
In Arabidopsis mutant myb98 the synergid cells are largely
normal but with abnormal FA pollen tube from wild type pollen
grow normally on the funiculus but fail to enter the micropyle.
2.One of the synergids forms the seat for pollen tube
discharge in the embryo sac.
3. Jensen(1965) suggested that the FA may be aiding the
synergid in the absorption and transportation of materials
into the embryo sac from the nucellus.FA forms the path of
entry for these substances.
• Many investigators have expressed doubts regarding the
nutritive role of the synergids.
• Current prevailing view is that the entry of metabolites
into the embryo sac is mainly through its chalazal end.
 Egg –
• Egg shows common walls with the two synergids
and the central cell. The wall is thicker in the
micropylar region but becomes thinner toward the
chalazal side.
• Young egg cell is richly endowed with organelles.At
maturity, however, organelles become scarce
indicating poor physiological activity.
 Antipodals –
• Usually they degenerate before or soon after
fertilization,without any appreciable enlargement.
• In sapotaceae(except Mimusops) and the Thismiaceae
the antipodal nuclei degenerate even without
organizing into cells.
• In many plants the antipodals are persistent and show
some structural and cytological features suggesting
their possible role in the nutrition of the embryo sac.
• In Caltha palustris they persist upto the octantstage of
the proembryo.
• Highest number of antipodal cells known is 300,recorded
in Sasa paniculata.
• Zea mays has about 20 antipodal cells,each with 1-4
nuclei. During additional divisions in antipodals, the
walls of many cells remain incomplete,leaving
protoplasmic continuties between adjacent cells.results
in the formation of multinucleate protoplasm or
syncytium.
• Antipodal nuclei may also become polyploid due to
endopolyploidy (Chrysanthemum) or polyteny (Papaver).
• Haustorial behaviour of antipodal cells is known in many plants
(Grindelia,Quinchamalium).
• In Quinchamalium chilense the antipodal nuclei do not organize
into individual cells. Instead they are cut off from the rest of
embryo sac by a wall,forming a multinucleate antipodal chamber.
The latter grows through the funiculus,reaching up to the tip of
the placenta where it branches profusely. The nuclei in the
antipodal haustorium become hypertrophied and may even divide
giving rise to 5-7 nuclei.
• Functions –
1. A nutritive role has been proposed for the antipodal cells,
especially where they are persistent. Show similarity with
other nutritive tissues,especially the glandular anther tapetum
and integumentary tapetum,in having increased DNA content
through multinucleate condition,endopolyploidy,or polyteny.
2. May also store large quantities of starch, lipids,and proteins
which are utilized by the developing endosperm and embryo.
3. Produce and secrete substances that control the growth and
development of endosperm.
 Central cell –
• Largest cell of the embryo sac, and the mother cell of the endosperm.
• Nuclei of the central cell, called polar nuclei.
• Two polar nuclei fuse,before or during double fertilization, to form
the secondary nucleus.
• Connected with the egg,synergids,and antipodals through
plasmodesmatal connections but no such connection exists between
the central cell and the adjacent nucellar cells.
• Presence of cell wall projections in the micropylar and/or chalazal
region shows that central cell draws nutrition from the surrounding
nucellus or integument.
 Naked Embryo Sac –
• In most of the angiosperms the embryo sac resides inside the
ovule,enclosed by the thick nucellus and the integuments.
• In some genera,such as Gallium,Philadelphus,Thesium,Torenia,
Utricularia and Vandellia,the embryo sac protrudes out through
the micropyle.
• In Torenia fournieri, the egg apparatus and the micropylar half of
the central cell are located outside the micropyle – easily
observed in intact ovules by differential interference
microscopy,making it a suitable material to study the entire
process of double fertilization.
• Extra –ovular extension of the embryo sac at the micropylar
end is quite common in santalaceae.
• In Loranthaceae the embryo sac grow upward up to various
heights. They reach the base of style in Macrosolen
cochinchinensis,and up to stigma in Helixanthera ligustrina.
• Longest embryo sacs occur in Moquiniella rubra. Here the tips
of the embryo sacs sometimes curve backward into the style by
2-4 mm after touching the stigma.
 Haustorial behaviour of embryo sac –
• In addition to the synergid and antipodal haustoria the entire embryo
sac may grow beyond the ovular tissue.
• It may also form micropylar or chalazal extensions or caeca from the
central cell which grow into various carpellary tissues and function as
haustoria.
• In Utricularia when the embryo sac is at the 2-nucleate stage the
nucellar epidermis and the remains of the non-functional megaspores
disappear completely.The tip of the embryo sac grows through the
ovule and comes in direct contact with the placenta.At the 4-nucleate
stage the embryo sac tip starts penetrating into the placental nutritive
tissue and at the mature embryo sac stage it is buried in the nutritive
tissue.
• Most of the Loranthaceae and santalaceous
members show the extension of the chalazal end
of the embryo sac in the form of a lateral
caecum,leaving the antipodal cells in situ.
Nutrition of embryo sac –
• Nucellus is the obvious pathway for nutrients to enter the
embryo sac because it surrounds the latter on all sides.
• The morphology of the ovule suggest that the main pathway of
nutrients into the embryo sac is the chalazal end.
• Funicular vascular supply,which can be regarded as the normal
channel for the supply of food materials to the ovule,
terminates at the base of the integuments.
• From there nutrients pass into the embryo sac through the
nucellar tissue present between the vascular supply and the
chalazal end of the embryo sac.
• Often hypostase is present in between the funicular vascular
supply and the chalazal end of the embryo sac. It has been
suggested by some authors that the hypostase may be involved in
the transfer of food materials to the embryo sac.
• While the entry of food material into the embryo sac appears to
be mainly through the chalazal end, the entire surface of the
female gametophyte seems to be absorbing nutrients from its
surrounding nucellar cells.