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The Questions of Developmental Biology

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Fig. 22.15<br />

Fig. 22.16<br />

<strong>The</strong> formation <strong>of</strong> the proximal-distal axis also proceeds similarly in vertebrates and<br />

invertebrates. In insects, the wing margin forms at the border between the dorsal cells and the<br />

ventral cells. <strong>The</strong> dorsal cells express Apterous protein, and this activates the expression <strong>of</strong><br />

Fringe. <strong>The</strong> interaction <strong>of</strong> the Fringe protein with the ventral cells leads to the growth <strong>of</strong> the wing<br />

blade outward from the body wall. Unexpectedly, a very similar cascade induces the outgrowth <strong>of</strong><br />

the vertebrate limb. <strong>The</strong> apical ectodermal ridge (AER) forms at the junction <strong>of</strong> the dorsal cells<br />

with the ventral cells. <strong>The</strong> dorsal cells express Radical fringe, a vertebrate homologue <strong>of</strong> the<br />

Fringe protein. This protein is critical in forming the AER. It is interesting that the ways in which<br />

these proteins become localized differ enormously between these phyla. Radical fringe, for<br />

instance, is induced by fibroblast growth factors, and it induces the AER to produce more<br />

fibroblast growth factors for the outgrowth <strong>of</strong> the limb. Fibroblast growth factors play no role in<br />

insect limb development. Yet, the instructions for limb outgrowth and polarity appear to be<br />

essentially the same in insects and vertebrates. Nature may have evolved the mechanism to form a<br />

limb only once, in the PDA, and both arthropods and vertebrates use modifications that process to<br />

this day.

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