The head problem. The organizational significance of segmentation in head development

This review argues for the segmental basis of chordate head organization which, like somite-based segmental organization in the trunk, takes its origin from early mesodermal development. The review builds on, and brings up to date, Goodrich's well-known scheme of head organization. It surveys recent data in support of this scheme and shows how evidence and arguments supposedly in conflict with it can be accommodated. Many of the arguments revolve around matters of methodology; the limitations of older LM, SEM (on which the concept of "somitomeres" is based) and recent molecular evidence (which has sometimes been seen as allocating the central role in head organization to the CNS and the neural crest) are highlighted and shown to explain a number of claims contrary to Goodrich's. We provide (in Part 2) a new, comparative survey of the best available evidence most directly relevant to the Goodrich Bauplan, with a special emphasis on stem chordates. The postotic region has commonly been seen as segmentally organized: the critical issues concern the preotic region. There are many reasons why Goodrich's three preotic segments may become specialized during evolution and why the underlying initial segmental organization may be overridden in later stages during embryonic development; we refer to a number of these. We conclude that the preotic segmental Bauplan is remarkably conserved and most explicitly demonstrated among stem forms, but we also suggest that the concept of the prechordal plate requires careful reexamination. Central to our overall analysis is the importance of the epigenetic nature of embryogenesis; its implications are made clear. Finally we speculate on evolutionary implications for the origin of the head and its specialized features. The review is intended to serve as a resource giving access to references to a wealth of now neglected, older data on anamniote embryology.