Somitogenesis in zebrafish

Both genetic and embryological studies in the zebrafish, Danio rerio, have contributed to our general understanding of how somites form and differentiate. In the zebrafish, mutants have been isolated that have specific effects on virtually every aspect of somite development. The fss-type mutants, defining 5 genes, affect somite segmentation and epithelialization. The you-type mutants, comprising 7 genes, and mutants in another 13 genes defective in notochord formation, have somites with abnormal pattern and morphology. Eighteen genes have been identified that are required for the differentiation and maintenance of the somitic musculature, and 2 genes have been identified that are involved in the development of motoneurons that innervate the somitic musculature. The true utility of the zebrafish lies in the ability to combine genetic analysis with embryological experimentation. Such analysis of somite segmentation suggests that homologues of both the Drosophila pair-rule and segment polarity genes, her1 and Sonic hedge-hog, respectively, are involved generating periodicity during somitogenesis. The Sonic hedge-hog protein secreted from the notochord also induces the formation of specific muscle types including the slow muscle fibers which are initially induced in the medial somite and undergo a series of morphological transitions including migration through the somite to the lateral surface where they complete their differentiation. The role of the notochord in patterning the somite is also demonstrated by its involvement in regulating the permissiveness of the somite to the extension of axons of primary motoneurons.