ZOO501 GDB Fall 2020 solution idea:
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 sperms 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such as sea urchins have a two-step polyspermy prevention strategy, with the fast, but transient, electrical block superseded after the first minute or so by a more slowly developing permanent mechanical block. It is thought that electrical blocks evolved in those 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.
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