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The Questions of Developmental Biology

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Lying immediately beneath the plasma membrane <strong>of</strong> the egg is a thin shell (about 5 μm)<br />

<strong>of</strong> gel-like cytoplasm called the cortex. <strong>The</strong> cytoplasm in this region is stiffer than the internal<br />

cytoplasm and contains high concentrations <strong>of</strong> globular actin molecules. During fertilization,<br />

these actin molecules polymerize to form long cables <strong>of</strong> actin known as micr<strong>of</strong>ilaments.<br />

Micr<strong>of</strong>ilaments are necessary for cell division, and they also are used to extend the egg surface<br />

into small projections called microvilli, which may aid sperm entry into the cell (see Figure 7.6B;<br />

also see Figure 7.19). Also within the cortex are the cortical granules (see Figures 7.4 and 7.6B).<br />

<strong>The</strong>se membrane-bound structures, which are homologous to the acrosomal vesicle <strong>of</strong> the sperm,<br />

are Golgi-derived organelles containing proteolytic enzymes. However, whereas each sperm<br />

contains one acrosomal vesicle, each sea urchin egg contains approximately 15,000 cortical<br />

granules. Moreover, in addition to digestive enzymes, the cortical granules contain<br />

mucopolysaccharides, adhesive glycoproteins, and hyalin protein. <strong>The</strong> enzymes and<br />

mucopolysaccharides are active in preventing other sperm from entering the egg after the first<br />

sperm has entered, and the hyalin and adhesive glycoproteins surround the early embryo and<br />

provide support for the cleavage-stage blastomeres.<br />

Many types <strong>of</strong> eggs also have an egg jelly outside the vitelline envelope (see Figure 7.4).<br />

This glycoprotein meshwork can have numerous functions, but most commonly is used either to<br />

attract or to activate sperm. <strong>The</strong> egg, then, is a cell specialized for receiving sperm and initiating<br />

development.<br />

*<strong>The</strong> contents <strong>of</strong> the egg vary greatly from species to species. <strong>The</strong> synthesis and placement <strong>of</strong> these materials will be<br />

addressed in Chapter 19, when we discuss the differentiation <strong>of</strong> germ cells<br />

Recognition <strong>of</strong> Egg and Sperm<br />

<strong>The</strong> interaction <strong>of</strong> sperm and egg generally proceeds according to five basic steps<br />

(Figure 7.8; Vacquier 1998):<br />

1. <strong>The</strong> chemoattraction <strong>of</strong> the sperm to the egg by soluble molecules secreted by the egg<br />

2. <strong>The</strong> exocytosis <strong>of</strong> the acrosomal vesicle to release its enzymes<br />

3. <strong>The</strong> binding <strong>of</strong> the sperm to the extracellular envelope (vitelline layer or zona pellucida) <strong>of</strong> the<br />

egg<br />

4. <strong>The</strong> passing <strong>of</strong> the sperm through this extracellular envelope<br />

5. Fusion <strong>of</strong> egg and sperm cell plasma membranes

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