Cause of polyspermy:

Physiological polyspermy happens when the egg normally accepts more than one sperm but only
one of the multiple sperm will fuse its nucleus with the nucleus of the egg. Physiological
polyspermy is present in some species of vertebrates and invertebrates.
Importance of prevention of polyspermy:
After successful fertilization of the egg cell, a block to polyspermy has to be established to avoid
sterility effects associated with lethal genome imbalance and chromosome segregation defects
during zygote and embryo development.

Two primary safeguards protect eggs against polyspermy:

● firstly, polyspermy is prevented at the plasma membrane level, where fusion of the first
sperm causes depolarization of the membrane and shedding of the egg’s sperm receptor
Folr4 (folate receptor 4; also known as Juno), thereby preventing fusion of further
sperm6,7.
● Secondly, fertilization triggers the exocytosis of cortical granules, a process termed
‘cortical reaction’. Cortical granules contain enzymes that modify and thereby harden the
zona pellucida, a proteinaceous matrix surrounding the oocyte8. This lowers the binding
affinity of sperm by cleavage of the zona pellucida protein ZP2 (ref. 9) and makes the
zona impermeable to additional sperm.

Fast block of polyspermy

The eggs of sexually-reproducing organisms are adapted to avoid this situation.The defenses are
particularly well characterized in the sea urchin, which respond to the acceptance of one sperm
by inhibiting the successful penetration of the egg by subsequent sperm. Similar defenses exist in
other eukaryotes.
The prevention of polyspermy in sea urchins depends on a change in the electrical charge across
the surface of the egg, which is caused by the fusion of the first sperm with the egg.Unfertilized
sea urchin eggs have a negative charge inside, but the charge becomes positive upon fertilization.
When sea urchin sperm encounter an egg with a positive charge, sperm-egg fusion is blocked.
Thus, after the first sperm contacts the egg and causes the change, subsequent sperm are
prevented from fusing. This "electrical polyspermy block" is thought to result because a
positively charged molecule in the sperm surface membrane is repelled by the positive charge at
the egg surface.
Electrical polyspermy blocks operate in many animal species, including frogs, clams, and marine
worms, but not in the several mammals that have been studied (hamster, rabbit, mouse). In
species without an electrical block, polyspermy is usually prevented by secretion of materials
that establish a mechanical barrier to polyspermy. Animals such as sea urchins have a two-step
polyspermy prevention strategy, with the fast, but transient, electrical block superseded after the
first minute or so by a more slowly developing permanent mechanical block. Electrical blocks
are helpful in species where a very fast block to polyspermy is needed, due to the presence of
many sperm arriving simultaneously at the egg surface, as occurs in animals such as sea urchins.
In sea urchins, fertilization occurs externally in the ocean, such that hundreds of sperm can
encounter the egg within several seconds.

Slow block of polyspermy

In mammals, in which fertilization occurs internally, fewer sperm reach the fertilization site in
the oviduct. This may be the result of the female genital tract being adapted to minimize the
number of sperm reaching the egg. Nevertheless, polyspermy preventing mechanisms are
essential in mammals; a secretion reaction, the "cortical reaction" modifies the extracellular coat
of the egg (the zona pellucida), and additional mechanisms that are not well understood modify
the egg's plasma membrane.The zona pellucida is modified by serine proteases that are released
from the cortical granules. The proteases destroy the protein link between the cell membrane and
the vitelline envelope, remove any receptors that other sperm have bound to, and help to form the
fertilization envelope from the cortical granules.
The cortical reaction occurs due to calcium oscillations inside the oocyte. What triggers such
oscillations is PLC-zeta, a phospholipase unique to sperm that is very sensitive to calcium
concentrations. When the first spermatozoa get inside the oocyte, it brings in PLC-zeta, that is
activated by oocyte's basal calcium concentrations, initiates the formation of IP3 and causes
calcium release from endoplasmic reticulum stores, generating the oscillations in calcium
concentration that will activate the oocyte and block polyspermy.