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Background  The importance of the cervical vertebrae as part of the skull–neck system in facilitating the success and diversity of tetrapods is clear. The reconstruction of its evolution, however, is problematic because of the variation in the number of vertebrae, making it difficult to identify homologous elements. Quantification of the morphological differentiation in the neck of diverse archosaurs established homologous units of vertebrae (i.e. modules) resulting from  Hox  gene expression patterns within the cervical vertebral column. The present study aims to investigate the modularity of the cervical vertebral column in the mouse and to reveal the genetic patterns and changes underlying the evolution of the neck of modern mammals and their extinct relatives. In contrast to modern mammals, non-mammalian synapsids are characterized by a variable cervical count, the presence of free cervical ribs and the presence of a separate CV1 centrum. How might these evolutionary modifications be associated with changes in the  Hox  code?    Results  In combination with up-to-date information on cervical  Hox  gene expression including description of the vertebral phenotype of  Hox  knock-out mutants, the 3D landmark-based geometric morphometric approach demonstrates a correlation between  Hox  code and vertebral morphology in the mouse. There is evidence that the modularity of the neck of the mouse had already been established in the last common ancestor of mammals, but differed from that of non-mammalian synapsids. The differences that likely occurred during the evolution of synapsids include an anterior shift in  HoxA-5  expression in relation to the reduction of cervical ribs and an anterior shift in  HoxD-4  expression linked to the development of the highly differentiated atlas-axis complex, whereas the remaining  Hox  genes may have displayed a pattern similar to that in mammals on the basis of the high level of conservatism in the axial skeleton of this lineage.    Conclusion  Thus, the mouse  Hox  code provides a model for understanding the evolutionary mechanisms responsible for the great morphological adaptability of the cervical vertebral column in Synapsida. However, more studies in non-model organisms are required to further elucidate the evolutionary role of  Hox  genes in axial patterning of the unique mammalian body plan.

Correlation between Hox code and vertebral morphology in the mouse: towards a universal model for Synapsida