On the homology of structures and Hox genes: the vertebral column

Research on expression patterns of Hox genes has revealed a surprisingly high conservation among vertebrates. In agreement with this conservation, a correlation has been found between the anterior limits of expression areas of certain Hox genes and the borders between morphological regions of the vertebral axis. These similarities are striking and important, but also counterintuitive, unless there are strong selection pressures to protect this conservatism. It is important to identify the selective forces that maintain these conservative networks. These selective forces can be due to pleiotropy or to internal selection. Discussed are the selective factors that are involved in the evolutionary constraint on the number of cervical vertebral numbers in mammals. Factors involved are due to internal selection and involve susceptibility to cancer, stillbirths and neuronal problems. It is intriguing how similar genetic networks can lead to fundamentally different animals. Clearly the same genes are used for different purposes. It is therefore important to try to find these differences. The search, for homology between organisms, and the enthusiasm about similarities that come with it, at times impedes the discovery of such differences. I have searched the literature for differences within vertebrates in the functioning and expression patterns of Hox genes during the development of the vertebral axis. The ensuing implications for homology of structures and genes are discussed. The vertebral column is a promising model system for the evaluation of the relationship between homologous Hox genes and homologous structures because of the large conservation of Hox gene expression patterns along the anterior axis. However, extensive remodelling of the vertebrate column indicates that important changes in the genetic basis must have taken place. A survey of the literature indicates that the correlation between Hox gene expression areas and vertebral regions is not such that one can predict the borders between vertebral regions on the basis of Hox gene expression patterns. The involvement of Hox genes in the development of identity of vertebrae is complex and the problems regarding the value of gene expression patterns for the determination of anatomical homology are discussed.