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The Questions of Developmental Biology

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Kessel and Gruss (1991) found this to be the case. Wild-type mice<br />

have 7 cervical (neck) vertebrae, 13 thoracic (ribbed) vertebrae, and 6<br />

lumbar (abdominal) vertebrae, in addition to the sacral and caudal (tail)<br />

vertebrae. In embryos exposed to RA on day 8 <strong>of</strong> gestation (during<br />

gastrulation), the first one or two lumbar vertebrae were transformed into<br />

thoracic (ribbed) vertebrae, while the first sacral vertebra <strong>of</strong>ten became a<br />

lumbar vertebra. In some cases, the entire posterior region <strong>of</strong> the mouse<br />

embryo failed to form (Figure 11.39E). <strong>The</strong>se changes in structure were<br />

correlated with changes in the constellation <strong>of</strong> Hox genes expressed in these tissues. For example,<br />

when RA was given to embryos on day 8, Hoxa-10 expression was shifted posteriorly, and an<br />

additional set <strong>of</strong> ribs formed on what had been the first lumbar vertebra. When posterior Hox<br />

genes were not expressed at all, the caudal part <strong>of</strong> the embryo failed to form.<br />

Retinoic acid probably plays a role in axis specification during normal development, and<br />

the source <strong>of</strong> retinoic acid is probably the node (Hogan et al. 1992; Maden et al. 1996). It is<br />

possible that the specification <strong>of</strong> mesoderm cells depends on the amount <strong>of</strong> time spent within the<br />

high retinoic acid concentrations <strong>of</strong> the node: the more time spent in the node, the more posterior<br />

the specification. This pattern can be demonstrated in culture, as embryonal carcinoma cells<br />

express more "posterior" Hox genes the longer they are exposed to retinoic acid (Simeone et al.<br />

1990). Giving RA exogenously would mimic the RA concentrations normally encountered only<br />

by the posterior cells. <strong>The</strong> evidence points to a Hox code wherein different constellations <strong>of</strong> Hox<br />

genes, activated by different retinoic acid concentrations, specify the regional characteristics<br />

along the anterior-posterior axis.<br />

Comparative anatomy.<br />

A new type <strong>of</strong> comparative embryology is now emerging, and it is based on the<br />

comparison <strong>of</strong> gene expression patterns. Gaunt (1994) and Burke and her collaborators (1995)<br />

have compared the vertebrae <strong>of</strong> the mouse and the chick. Although the mouse and the chick have<br />

a similar number <strong>of</strong> vertebrae, they apportion them differently. Mice (like all mammals, be they<br />

giraffes or whales) have only 7 cervical vertebrae. <strong>The</strong>se are followed by 13 thoracic vertebrae, 6<br />

lumbar vertebrae, 4 sacral vertebrae, and a variable (20+) number <strong>of</strong> caudal vertebrae (Figure<br />

11.40).

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