Abstract
BackgroundThe 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?ResultsIn 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.ConclusionThus, 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.
Highlights
The importance of the cervical vertebrae as part of the skull–neck system in facilitating the success and diversity of tetrapods is clear
About 84% of the total variance in the sample is explained by the first two relative warps (RW) (Table 2) and, the morphospace constructed from RW 1 and RW 2 provides a reasonable approximation of the total shape variation (Fig. 2)
The scatter plot shows that the axis (CV2) is in the second quadrant, the three cervical vertebrae (CV3-5) are in the fourth quadrant, whereas the last two cervical vertebrae (CV6-7) are in the first quadrant
Summary
The importance of the cervical vertebrae as part of the skull–neck system in facilitating the success and diversity of tetrapods is clear. 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. 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?. The evolution of a morphologically distinct and functional neck consisting of a series of cervical vertebrae (CV) had a great impact on the ecological diversification of tetrapods due to its involvement in a number of vital functions [1,2,3]. Even within the cervical vertebral column of tetrapods, differences in the number of vertebrae and, in the morphological regionalization of the neck correspond to modifications in Hox gene expression domains (expansion of a Hox gene’s expression domain and/or a shift of gene expression) [13]
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