Fertilization

• The fusion of male and female gametes during

sexual reproduction to form a diploid zygote is
called fertilization.
• Process of fertilization was first discovered by
Strasburger in Monotropa (1884).
• In angiosperms, the male gamete could not
directly fuse with female gamete just after
pollination since the female gamete lies deep
in the ovarian cavity.
• For the fertilization, pollen grains germinate in
the stigma and develop pollen tube.
• The pollen tube grows and reaches to embryo
sac to discharge two male gametes near the
egg.
• Out of two gametes, one fuses with the egg
(forming zygote) and other fuses with the
polar nuclei or secondary nucleus to form
primary endosperm nucleus.
• The phenomenon in which the male gametes
are brought to the female gametophyte by a
pollen tube is called siphonogamy.
 The process of fertilization involves:

1. Germination of pollen grains:

2. Growth of pollen tube
3. Entry of pollen tube into the ovule:
4. Entry of pollen tube into embryo sac
5. Discharge of male gametes from pollen tube
6. Syngamy (true fertilization)
7. Triple fusion:
 1. Germination of pollen grains:
• After pollination, the pollen grains are deposited on the
stigma where they germinate and produce pollen tube
(which represents the male gametophytes of
angiosperms)
• time taken for the pollen germination on stigma may vary
from few minutes to several days, according to the
species.
• Usually one pollen tube comes out from each pollen grain
(monosiphonous) but in some members of Malvaceae,
Cucurbitaceae, Campanulaceae etc.several pollen tubes
(10-14) have been reported developing from the same
pollen grain (Polysiphonous) but only one of them finally
matures, makes further progress and becomes functional.
• In the first step of the germination, pollen grain
absorbs the stigmatic fluid or sugar solution,
gets hydrated and swells up.
• After hydration, swelling of (outer) pectic layer
of intine occurs which leads to the rupture of
thin exine layer in germ pore region.
• Then the intine protrudes out through the germ
pore giving small tubular out growth called
pollen tube.
• Almost the entire content of the pollen grain
moves into the tube.
• In a growing tube, most of the cytoplasm is
confined to the apical region and a large vacuole
fills the grain and older region of the tube.
• Cytoplasm is restricted to the apical region by
the formation of series of callose plugs behind
the tip as result fully grown pollen tube is sub—
divided into many compartments.
• Partition in the pollen tube prevents the
backflow of cytoplasm and nuclei.
• Organization of the pollen tube is achieved after it has
attained a considerable length and the generative cell
and vegetative nucleus have moved into it and shows
four distinct zones:
• i) Apical zone/growth region: swollen region, contains
abundant vesicles, lacks cell organelles.
• ii) Sub-apical zone: Have granular cytoplasm rich in
cell organelles, vesicles are produced in this region.
• iii) Nuclear zone: Consists vegetative nucleus and
generative cell (later two sperms)
• iv) Vacuolization zone: forms a transition between the
active and inactive cytoplasm of the pollen tube, it is
highly vacuolated and ends with a callosic plug.
Organization of pollen tube
 2. Growth of pollen tube:
• The rapid growth of the pollen tube is restricted to the
tip region.
• The pollen tube further grows and passes through the
stigma and style to the ovary.
• Style may be solid or semi-solid in dicot and open in
monocot.
• In case of solid style, it is filled with the transmitting
tissues or conducting tissue consisting of 2-3 layers of
elongated glandular cells and in hollow style there is a
canal which runs through out stigma to the base of style.

 
• The canal is lined by a single layer of glandular cells
which secrete some mucilaginous substance.
• The pollen tube secretes some enzymes which
dissolve the tissues of style making its way to the
ovary.
• vegetative nucleus (tube nucleus) located near the
growing tip regulates the growth of the pollen tube.
• The generative nucleus divides to form two male
nuclei which soon form male gametes.
 3. Entry of pollen tube into the ovule:
• After reaching ovary, the pollen tube enters into the ovule
through micropyle and sometimes through other regions
(chalaza, integument). Based on the path of entry of
pollen tube into the ovule, fertilization is of three types:
• Porogamy: The pollen tube enters into the ovule through
micropyle, most common type. Eg: lily.
• Mesogamy: The pollen tube enters into the ovule through
integuments (Cucurbita) or through Funiculus (Pistacia)
• Chalazogamy: The pollen tube enters into the ovule
through chalaza. E.g. Casuarina
• With respect to the porogamy, some worker
suggests that the entry of pollen tube into the
ovule and its further growth towards the embryo
sac is regulated by a chemotropic substances
secreted by filiform apparatus of synergids into the
micropyle.
• According to Rosen (1965) the chemotropic
substance is secreted by micropyle itself rather
than the synergids.
• Chao (1972) demonstrated that the distal part of
the integuments secrets mucilaginous substance
into the micropyle by its dissolution, which directs
the path of pollen tube and help in its growth both
mechanically and chemotropically.
• A special structure called obturator (comprising
multicellular glandular epidermal hairs
originating from the placenta or Funiculus)
found in some plants, forms a kind of bridge for
the pollen tube to reach the ovule.
• Their surface exudates provide nutrition and
chemical guidance to the pollen tube.
• After fertilization, the obturator shrinks and
disappears.
 4. Entry of pollen tube into embryo sac:
• After entering the ovules either through the
micropyle or through chalaza or by piercing
the integuments or Funiculus, the pollen tube
passes through the nucellus and ultimately
penetrates the wall of embryosac through the
micropylar end.
• The pollen tube may either pass between the
egg and one synergid or between the embryo
sac wall and a synergid or directly into the
synergid.
• When it enters through one of the synergids, it
gets degenerated by the impact of pollen tube
and the other remains intact for some period of
time.
• Later the remaining synergid also gets
degenerated.
• In some case both synergids degenerate before
the entry of pollen tube into the embryo sac.
5. Discharge of male gametes from pollen tube:

• Ultimately the tip of the pollen tube bursts

and both the male gametes are discharged
into the embryo sacs.
• The tube nucleus is generally disorganized by
this time.
 6. Syngamy (true fertilization):

• Out of the two male gametes, one fuses with

the egg nucleus and forms the diploid zygote.
• This process of gametic fusion is called
syngamy or true fertilization.
• The zygote on its further development forms
the embryo.
 7. Triple fusion:
• The second male gamete fuses with the two
haploid polar nuclei or diploid secondary
nucleus to form a triploid nucleus or primary
endosperm nucleus.
• This process involving the fusion of three
nuclei is called triple fusion.
• The primary endosperm nucleus gives rise to
the endosperm.
 Double fertilization
• The phenomenon of fertilization involving the
fusion of one of the male gamete with the egg
together with the fusion of second male gamete
with the polar nuclei is called double fertilization.
• It involves both syngamy and triple fusion.
• Double fertilization is a unique feature of
angiosperm and is necessary for the production
of viable seeds in the flowering plants.
• It was first observed by Nawaschine(1898) in
Fritillaria and Lillium.
 Significance of double fertilization

 
1. It is necessary for the production of viable
seeds. The embryo formed after true fertilization
gives rise to baby plant and the endosperm
formed after the triple fusion develops into the
endosperm which provides the nourishment for
the developing embryo.
 
2. In angiosperms the growth of the embryo sac
(female gametophyte) stops at 8 nucleate or 7
celled stages. The fusion of second male gamete
with the polar nuclei or secondary nucleus
provides stimulus to resume growth and form the
nutritive tissue or endosperm from the secondary
nucleus.
i.e. without double fertilization the secondary
nucleus would have remained inactive and no
endospermic tissue would have been formed.
3. Double fertilization ensures that the nutritive tissue
or endosperm is formed only when the formation of
embryo has been confirmed by fertilization of egg. If
fertilization fails due to some reason, no endosperm
will be formed. Thus no energy of the plant will be
wasted on this account in angiosperms.

4. The endospermic nuclei have maternal and

paternal chromosomes thus the endospermic tissue
shows the physiological aggressiveness due to the
hybrid vigor.