EGG ACTIVATION AT FERTILIZATION BY A SOLUBLE SPERM PROTEIN

The most fundamental unresolved issue of fertilization is to define how the sperm activates the egg to begin embryo development. Egg activation at fertilization in all species thus far examined is caused by some form of transient increase in the cytoplasmic free Ca2+ concentration. What has not been clear, however, is precisely how the sperm triggers the large changes in Ca2+ observed within the egg cytoplasm. Here, we review the studies indicating that the fertilizing sperm stimulates a cytosolic Ca2+ increase in the egg specifically by delivering a soluble factor that diffuses into the cytosolic space of the egg upon gamete membrane fusion. Evidence is primarily considered in species of eggs where the sperm has been shown to elicit a cytosolic Ca2+ increase by initiating Ca2+ release from intracellular Ca2+ stores. We suggest that our best understanding of these signaling events is in mammals, where the sperm triggers a prolonged series of intracellular Ca2+ oscillations. The strongest empirical studies to date suggest that mammalian sperm-triggered Ca2+ oscillations are caused by the introduction of a sperm-specific protein, called phospholipase C-zeta (PLC zeta) that generates inositol trisphosphate within the egg. We will discuss the role and mechanism of action of PLC zeta in detail at a molecular and cellular level. We will also consider some of the evidence that a soluble sperm protein might be involved in egg activation in nonmammalian species.