Developmental control of segment numbers in vertebrates.

Segmentation or metamery in vertebrates is best illustrated by the repetition of the vertebrae and ribs, their associated skeletal muscles and blood vessels, and the spinal nerves and ganglia. The segment number varies tremendously among the different vertebrate species, ranging from as few as six vertebrae in some frogs to as many as several hundred in some snakes and fish. In vertebrates, metameric segments or somites form sequentially during body axis formation. This results in the embryonic axis becoming entirely segmented into metameric units from the level of the otic vesicle almost to the very tip of the tail. The total segment number mostly depends on two parameters: (1) the control of the posterior growth of the body axis during somitogenesis-more same-size segments can be formed in a longer axis and (2) segment size--more smaller--size segments can be formed in a same-size body axis. During evolution, independent variations of these parameters could explain the huge diversity in segment numbers observed among vertebrate species. These variations in segment numbers are accompanied by diversity in the regionalization of the vertebral column. For example, amniotes can exhibit up to five different types of vertebrae: cervical, thoracic, lumbar, sacral and caudal, the number of which varies according to the species. This regionalization of the vertebral column is controlled by the Hox family of transcription factors. We propose that during development, dissociation of the Hox- and segmentation-clock-dependent vertebral patterning systems explains the enormous diversity of vertebral formulae observed in vertebrates.