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Background   Hox  genes play a central role in axial patterning during animal development. They are clustered in the genome and specify cell fate in sequential domains along the anteroposterior (A-P) body axis in a conserved order that is co-linear with their relative genomic position. In the soil worm  Caenorhabditis elegans , this striking rule of co-linearity is broken by the anterior  Hox  gene  ceh-13 , which is located between the two middle  Hox  paralogs,  lin-39  and  mab-5 , within the loosely organized nematode  Hox  cluster. Despite its evolutionary and developmental significance, the functional consequence of this unusual genomic organization remains unresolved.    Results  In this study we have investigated the role of  ceh-13  in different developmental processes, and found that its expression and function are not restricted to the anterior body part. We show that  ceh-13  affects cell migration and fusion as well as tissue patterning in the middle and posterior body regions too. These data reveal novel roles for  ceh-13  in developmental processes known to be under the control of middle  Hox  paralogs. Consistently, enhanced activity of  lin-39  and  mab-5  can suppress developmental arrest and morphologic malformation in  ceh-13  deficient animals.    Conclusion  Our findings presented here show that, unlike other  Hox  genes in  C. elegans  which display region-specific accumulation and function along the A-P axis, the expression and functional domain of the anterior  Hox  paralog  ceh-13  extends beyond the anterior region of the worm. Furthermore,  ceh-13  and the middle  Hox  paralogs share several developmental functions. Together, these results suggest the emergence of the middle-group  Hox  genes from a  ceh-13 -like primordial  Hox  ancestor.

The C. elegans Hox gene ceh-13 regulates cell migration and fusion in a non-colinear way. Implications for the early evolution of Hoxclusters