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Pollen-Pistil Interaction: Pollen Attachment and Hydration

Pollen grains must successfully interact with the pistil in order to fertilize the female gametes. When pollen lands on the stigma, it must hydrate, germinate, and form a pollen tube to deliver sperm cells through the style tissue. The pistil provides nutrients and signals to guide compatible pollen tubes specifically to female gametes. This pollen-pistil interaction is crucial for angiosperm reproduction and allows for selection of viable pollen through competition during growth.

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1K views

Pollen-Pistil Interaction: Pollen Attachment and Hydration

Pollen grains must successfully interact with the pistil in order to fertilize the female gametes. When pollen lands on the stigma, it must hydrate, germinate, and form a pollen tube to deliver sperm cells through the style tissue. The pistil provides nutrients and signals to guide compatible pollen tubes specifically to female gametes. This pollen-pistil interaction is crucial for angiosperm reproduction and allows for selection of viable pollen through competition during growth.

Uploaded by

Sreeja Raj
Copyright
© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
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Download as DOCX, PDF, TXT or read online on Scribd
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Pollen-Pistil Interaction

Pollen grains are deposited on the stigma either due to closeness of


the anthers to the stigma or by pollinating agents (biotic or abiotic).
This unique feature brings about pollen-pistil interaction between
the male gametophyte, the pollen grains, with the massive
sporophytic tissue.

Pollen Attachment and Hydration:

The attachment of the pollen on the stigma depends upon its wall
sculpture and stickiness. In wet stigma adhesion is mostly a
mechanical process, whereas, in dry stigma it depends on the extent
and composition of the pellicle, and the amount of surface-coat
substances on the pollen.

Pollen hydration proceeds in a controlled manner characterized by


distinct area of stability of increasing water content and can begin
in the anther before pollen release.

Pollen Germination and Tube Growth:

The stigmatic surface provides the essential prerequisites for a


successful germination that are absent in the pollen. In wet stigma,
the role of the stigmatic exudates in pollen germination is highly
variable.

In dry stigma, the pellicle plays a vital role in germination. Its


enzymatic removal inhibits pollen germination or pollen tube entry
into stigma. For instance in Raphanus sativus enzymatic digestion
of the pellicle reduces pollen germination and totally inhibits the
entry of even compatible tubes into the stigmatic papillae.
The stigmatic surface also provides boron and calcium which are
required for germination but are deficient in pollen. It has been
seen that those stigmatic secretion of Vitis vinifera that contain 2-5
ppm of boron permit pollen germination.

The growth of the pollen tube of flowering plants is restricted


exclusively at their apices. Microscopic examination of growing
pollen tubes reveals that most of the cytoplasm is restricted to the
apical region while a large vacuole fills the grain and the older
region of the tube (Figure 6.6). The cytoplasm is restricted to the
apical region of the growing tube by the formation of series of
callose plugs at regular intervals behind the tip

In wet stigma and solid style the cuticle gets disrupted during the
secretion of the exudates and the pollen tube enters the intercellular
matrix of the stigmatic tissue.

Path of Pollen Tube in the Style:

In species with wet stigma and solid style the cuticle of the stigma
/papillae is disrupted during the secretion of the exudates, thus
there is no physical barrier for pollen tube entry into the
intercellular spaces of the transmitting tissue of the stigma. In taxa
with wet stigma and hollow style, pollen tubes grow on the surface
of the stigma and enter the stylar canal.

In species with dry stigma and solid style the cuticle provides the
physical barrier for the pollen tube entry. The cuticle is eroded at
the point of contact by the activity of cutinases released by the
pollen. After the digestion of the cuticle, the tube enters the
pectocellulosic wall of the papillae and finally grows through the
intercellular substances of the stigma and the style.

Path of Pollen Tube in the Ovule and Embryo Sac:

The pollen tube finally pushes through the ovule and reaches the
embryo sac. This guidance into the ovule is in terms of essential
signals originating from the male and female tissues.
Evidences obtained from the analysis of developmental mutants of
Arabidopsis, viz., bell and sinl (where integument and embryo sac
development in the ovule is affected) suggests that genes active in
the female gametophyte play a crucial role in the signalling process
that guides pollen tubes to the ovule.

In fact pollen tubes are always attracted to ovules with a functional


embryo sac, which confirms a female gametophytic control of pollen
tube guidance.

Recognition of the Pollen by the Stigma:

The stigmatic surface of a flower provides refuge to various pollen


grains, but a physiological mechanism operates to ensure that only
intraspecific pollen germinate successfully. In sporophytic self-
incompatibility system the recognition reaction system sets in
almost immediately after the pollen comes in contact with the
stigma.

This is also true for certain gametophytic self- incompatibility


systems. The components of pollen recognition system are present
in all floral organs, but are segregated to the surface of the stigma by
the action of genes like PDH.

Recognition of compatible pollen grains by the stigmatic papillae


involves a molecular interaction between substances present in the
pollen wall and those present on the pellicle or in the stigmatic
surface.

In fact recognition mechanism is switched on with the hydration of


pollen and the subsequent release of its wall proteins. The
molecular events that are switched on, following the acceptance or
rejection of the pollen grains have been studied in Brassica napus,
B. oleracea, and Arabidopsis, from where the details are gradually
emerging out.

The pollen grains following contact with the stigma synthesis nearly
forty new proteins, and few of these proteins are highly
phosphorylated. Thus it appears that protein phosphorylation may
be responsible for signal transduction in a compatible association.
Moreover, there is a brief Ca2+ peaks in the stigmatic papillae of
Brassica napus following its contact with compatible pollen grains.
Thus there is a definite participation of a Ca2+ in pollen signal
perception but how the system is activated to promote hydration
and germination of pollen grains is not clear.

The involvement of a specific component of the pollen-wall-based


tryphine in the regulation of pollen-stigma interactions has been
provided by the mutational study in Arabidopsis. Preuss (1993)
isolated a mutant (pop1) Arabidopsis, which was defective in pollen-
pistil interactions, thus inducing male sterility.

The grains failed to hydrate on the stigma and germinate, though it


responded successfully in vitro. The failure of the mutant to
germinate in vivo was possibly due to loss of tryphine components
on the wall of mutant pollen grains, and this was substantiated by
the absence of wax on the stem of pop1 mutants.

Further chemical analysis has shown that wax deficiency on the


stem is correlated with the absence of long chain lipids on the
mutant pollen grains. Further a mutation in the CER1 locus of
Arabidopsis which affect pollen hydration on the stigma, have also
been traced to lack functional tryphine layer with the normal
components of lipids on the pollen grains.

The gene has also been shown to encode for a new protein, involved
in catalytic steps of wax biosynthesis. Thus lipid molecules and
tryphine are involved in the signaling mechanism that allows the
stigma to have a successful pollination event.

Pollen grains of a transgenic Petunia hydrida plant, deficient in


flavonoids failed to germinate or produced very short pollen tube in
vitro. This inhibition was nullified by the addition of stigma extract
from the wild type Petunia or by adding micromolar quantities of
flavonoids (kaempferol) to the germinating medium.

Sharma and Shivanna (1983 a, b) investigated the biochemical basis


of self- incompatibility recognition in Petunia hybrida by using in
vitro assay. Pollen grains were cultured in a medium containing
pistil extract and before that the grains were treated with sugars or
with lectins.

Interestingly self-incompatibility could be avoided when the pollen


grains were treated with glucose or lactose but not with lectins thus
indicating that lectin-like components are involved in self-
incompatibility recognition. Blocking these molecules with
corresponding sugars make them ineffective in establishing
recognition, and thus overcome inhibition.

Likewise, inclusion of specific lectins into medium containing pistil


extract was also effective in overcoming inhibition of self-pollen.
Seemingly, recognition factors in the pistil are polysaccharide-
containing molecules and the binding of the saccharides with the
specific lectins makes them ineffective in recognizing self-pollen.

Significance of Pollen Pistil Interaction:

Pollen-pistil interaction is unique in flowering plants and plays a


significant role in sexual reproduction and seed formation. The
pollen grains carrying the male gametes do not have a direct
admittance to the female gamete.

Thus they need to be deposited on the stigma and it’s pollen tube
has to grow through the massive sporophytic tissues of the stigma
and style prior to release of male gametes near the egg. The pistil
has developed a unique mechanism to recognize pollen grains and
thereby permit the growth of pollen tube in compatible cases.

Incompatible tube is thus effectively prevented from reaching the


embryo sac. The barrier to fertilization is restricted to the
sporophytic tissues of the pistil. At the same time there is no chance
of unsuccessful fertilization ones the male gametes have been
released near the egg.

The principal significance of pollen-pistil interactions are:

a. The most essential requirement for sexual reproduction is the


screening and selection of male gamete and this is achieved during
pollen pistil interaction. Thus, pollen-pistil interaction offers
enormous potential for the manipulation of pollen screening which
is obviously for the quality and compatibility of pollen.

b. The number of pollen grains that are generally deposited on the


stigma under normal conditions are far greater than the number of
ovules available for fertilization. As a result the pollen grains are
subjected to a tough competition during pollen-pistil interaction.

Only those pollen that germinate early and have a faster growing
pollen tube, i.e., more vigorous, are able to withstand the rigidity of
post- pollination competition and fertilization. Consequently
competition among pollen grains during pollen-pistil interaction
results in the increased vigour of the progeny. Thus this interaction
can be considered as an important contributory factor in the
evolutionary success of flowering plants.

c. It has a direct relevance to plant breeding programmes. A plant


breeder continuously strives to bring together desirable characters
present in different taxa, through hybridization.

Thus a better understanding of the biology of pollen-pistil


interaction would no doubt, enable the plant breeders to
manipulate the screening process in the pistil more effectively.

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