Cortical Reaction in Sea Urchin | Slow Blocks to Polyspermy

The cortical reaction is a process initiated during fertilization by the release of cortical granules from the egg, which prevents polyspermy, the fusion of multiple sperm with one egg. In contrast to the fast block of polyspermy which immediately but temporarily blocks additional sperm from fertilizing the egg, the cortical reaction gradually establishes a permanent barrier to sperm entry and functions as the main part of the slow block of polyspermy in many animals.

To create this barrier cortical granules, specialized secretory vesicles located within the egg's cortex (the region directly below the plasma membrane), are fused with the egg's plasma membrane, causing the contents of the cortical granules to be released outside the cell where they modify an existing extracellular matrix to make it impenetrable to sperm entry. The cortical granules contain proteases that clip perivitelline tether proteins, peroxidases that harden the vitelline envelope, and glycosaminoglycans that attract water into the perivitelline space, causing it to expand and form the hyaline layer. The trigger for the cortical granules to exocytose is the release of calcium ions from cortical smooth endoplasmic reticulum in response to sperm binding to the egg.

In the well-studied sea urchin model system, the granule contents modify a protein coat on the outside of the plasma membrane (the vitelline layer) so that it is released from the membrane. The released cortical granule proteins exert a colloid osmotic pressure causing water to enter the space between the plasma membrane and the vitelline layer, and the vitelline layer expands away from the egg surface. This is easily visible through a microscope and is known as "elevation of the fertilization envelope". Some of the former granule contents adhere to the fertilization envelope, and it is extensively modified and cross-linked. As the fertilization envelope elevates, non-fertilizing sperm are lifted away from the egg plasma membrane, and as they are not able to pass through the fertilization envelope, they are prevented from entering the egg. Therefore, the cortical reaction prevents polyspermic fertilization, a lethal event. Another cortical granule component, polysaccharide-rich hyalin, remains adherent to the outer surface of the plasma membrane, and becomes part of the hyaline layer.