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Transposition of  Hoxd  genes to a more posterior (5′) location within the  HoxD  complex suggested that colinearity in the expression of these genes was due, in part, to the existence of a silencing mechanism originating at the 5′ end of the cluster and extending towards the 3′ direction. To assess the strength and specificity of this repression, as well as to challenge available models on colinearity, we inserted a  Hoxb1/lacZ transgene within the posterior  HoxD  complex, thereby reconstructing a cluster with a copy of the most anterior gene inserted at the most posterior position. Analysis of  Hoxb1  expression after ectopic relocation revealed that  Hoxb1 -specific activity in the fourth rhombomere was totally abolished. Treatment with retinoic acid, or subsequent relocations toward more 3′ positions in the HoxD  complex, did not release this silencing in hindbrain cells. In contrast, however, early and anterior transgene expression in the mesoderm was unexpectedly not suppressed. Furthermore, the transgene induced a transient ectopic activation of the neighboring Hoxd13  gene, without affecting other genes of the complex. Such a local and transient break in colinearity was also observed after transposition of the  Hoxd9/lacZ  reporter gene, indicating that it may be a general property of these transgenes when transposed at an ectopic location. These results are discussed in the context of existing models, which account for colinear activation of vertebrate  Hox  genes.

Mechanisms of Hox gene colinearity: transposition of the anterior Hoxb1 gene into the posterior HoxD complex