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The Questions of Developmental Biology

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Within this enormous volume <strong>of</strong> cytoplasm resides a large nucleus. In some species (e.g.,<br />

sea urchins), the nucleus is already haploid at the time <strong>of</strong> fertilization. In other species (including<br />

many worms and most mammals), the egg nucleus is still diploid, and the sperm enters before the<br />

meiotic divisions are completed. <strong>The</strong> stage <strong>of</strong> the egg nucleus at the time <strong>of</strong> sperm entry in<br />

different species is illustrated in Figure 7.5.<br />

Enclosing the cytoplasm is the egg plasma membrane. This membrane must regulate the<br />

flow <strong>of</strong> certain ions during fertilization and must be capable <strong>of</strong> fusing with the sperm plasma<br />

membrane. Outside the plasma membrane is the vitelline envelope (Figure 7.6), which forms a<br />

fibrous mat around the egg. This envelope contains at least eight different glycoproteins and is<br />

<strong>of</strong>ten involved in sperm-egg recognition (Correia and Carroll 1997). It is supplemented by<br />

extensions <strong>of</strong> membrane glycoproteins from the plasma membrane and by proteinaceous vitelline<br />

posts that adhere the vitelline envelope to the membrane (Mozingo and Chandler 1991).<br />

<strong>The</strong> vitelline envelope is essential for the species-specific binding <strong>of</strong> sperm. In mammals, the<br />

vitelline envelope is a separate and thick extracellular matrix called the zona pellucida.<br />

<strong>The</strong> mammalian egg is also surrounded by a layer <strong>of</strong> cells called the cumulus (Figure 7.7), which<br />

is made up <strong>of</strong> the<br />

ovarian follicular cells<br />

that were nurturing the<br />

egg at the time <strong>of</strong> its<br />

release from the ovary.<br />

Mammalian sperm<br />

have to get past these<br />

cells to fertilize the<br />

egg. <strong>The</strong> innermost<br />

layer <strong>of</strong> cumulus cells,<br />

immediately adjacent<br />

to the zona pellucida,<br />

is called the corona radiata.

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