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The Questions of Developmental Biology

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On their lateral sides, each cell has another cell for a neighbor. Fink and McClay found that the<br />

prospective ectoderm and endoderm cells (descendants <strong>of</strong> the mesomeres and macromeres,<br />

respectively) bind tightly to one another and to the hyaline layer, but adhere only loosely to the<br />

basal lamina (Table 8.2).<br />

Table 8.2. Affinities <strong>of</strong> mesenchymal and nonmesenchymal cells to cellular and extracellular<br />

components a<br />

Dislodgment force (in dynes)<br />

Cell type Hyaline Gastrula cell monolayers Basal lamina<br />

16-cell-stage micromeres 5.8 × 10 -5 6.8 × 10 -5 4.8 × 10 -7<br />

Migratory-stage mesenchyme cells 1.2 × 10 -7 1.2 × 10 -7 1.5 × 10 -5<br />

Gastrula ectoderm and endoderm 5.0 × 10 -5 5.0 × 10 -5 5.0 × 10 -7<br />

Source: After Fink and McClay 1985.<br />

a Tested cells were allowed to adhere to plates containing hyaline, extracellular basal lamina, or cell monolayers. <strong>The</strong><br />

plates were inverted and centrifuged at various strengths to dislodge the cells. <strong>The</strong> dislodgement force is calculated<br />

from the centrifugal force needed to remove the test cells from the substrate.<br />

<strong>The</strong> micromeres originally display a similar pattern <strong>of</strong> binding. However, the micromere<br />

pattern changes at gastrulation. Whereas the other cells retain their tight binding to the hyaline<br />

layer and to their neighbors, the primary mesenchyme precursors lose their affinity for these<br />

structures (which drops to about 2% <strong>of</strong> its original value) while their affinity for components <strong>of</strong><br />

the basal lamina and extracellular matrix (such as fibronectin) increases a hundredfold. This<br />

change in affinity causes the micromeres to release their attachments to the external hyaline layer<br />

and their neighboring cells and, drawn in by the basal lamina, migrate up into the blastocoel<br />

(Figures 8.18, 8.19). <strong>The</strong>se changes in affinity have been correlated with changes in cell surface<br />

molecules that occur during this time (Wessel and McClay 1985).<br />

As shown in Figure 8.18, there is a heavy concentration <strong>of</strong> extracellular material around<br />

the ingressing primary mesenchyme cells (Galileo and Morrill 1985; Cherr et al. 1992). Once<br />

inside the blastocoel, the primary mesenchyme cells appear to migrate along the extracellular<br />

matrix <strong>of</strong> the blastocoel wall, extending their filopodia in front <strong>of</strong> them (Galileo and Morrill 1985;<br />

Karp and Solursh 1985). Several proteins (including fibronectin and a particular sulfated<br />

glycoprotein) are necessary to initiate and maintain this migration (Wessel et al. 1984; Sugiyama<br />

1972; Lane and Solursh 1991; Berg et al. 1996).

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