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The Questions of Developmental Biology

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Ingression <strong>of</strong> primary mesenchyme<br />

Function <strong>of</strong> primary mesenchyme cells<br />

Shortly after the blastula hatches from<br />

its fertilization envelope, the vegetal side <strong>of</strong> the<br />

spherical blastula begins to thicken and flatten<br />

(Figure 8.17, 9 hours). At the center <strong>of</strong> this flat<br />

vegetal plate, a cluster <strong>of</strong> small cells begins to<br />

change. <strong>The</strong>se cells begin extending and<br />

contracting long, thin (30 × 5 μm) processes<br />

called filopodia from their inner surfaces. <strong>The</strong><br />

cells then dissociate from the epithelial<br />

monolayer and ingress into the blastocoel<br />

(Figure 8.17, 9 10 hours). <strong>The</strong>se cells, derived<br />

from the micromeres, are called the primary<br />

mesenchyme. <strong>The</strong>y will form the larval<br />

skeleton, so they are sometimes called the<br />

skeletogenic mesenchyme.<br />

At first the cells appear to move randomly along<br />

the inner blastocoel surface, actively making and<br />

breaking filopodial connections to the wall <strong>of</strong><br />

the blastocoel. Eventually, however, they<br />

become localized within the prospective<br />

ventrolateral region <strong>of</strong> the blastocoel. Here they<br />

fuse into syncytial cables, which will form the<br />

axis <strong>of</strong> the calcium carbonate spicules <strong>of</strong> the<br />

larval skeleton (Figure 8.18).<br />

Importance <strong>of</strong> extracellular lamina inside the blastocoel<br />

<strong>The</strong> ingression <strong>of</strong> the micromere descendants into the blastocoel<br />

is caused by these cells losing their affinity for their neighbors and for<br />

the hyaline membrane and acquiring a strong affinity for a group <strong>of</strong><br />

proteins that line the blastocoel. This model was first proposed by<br />

Gustafson and Wolpert (1967) and was confirmed in 1985, when Rachel<br />

Fink and David McClay measured the strengths <strong>of</strong> sea urchin blastomere<br />

adhesion to the hyaline layer, to the basal lamina lining the blastocoel,<br />

and to other blastomeres. Originally, all the cells <strong>of</strong> the blastula are<br />

connected on their outer surface to the hyaline layer and on their inner<br />

surface to a basal lamina secreted by the cells.

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