Clarifying tetrapod embryogenesis, a physicist's point of view

The origin of tetrapods is a complex question that webs together genetic, paleontological, developmental and physical facts. Basically, the development of embryos is described by a complex mix of mechanical movements and biochemical inductions of genetic origin. It is difficult to sort out in this scientific question what are the fundamental features imposed by conservation laws of physics, and by force equilibria, and what can be ascribed to successive, very specific, stop-and-go inductions of genetic nature. A posteriori, evolution selects the parameters of this process as found in the observed species. Whether there is a general law to animal formation seems out of the question. However, several concepts developed in biology, like the concept of organizer seem questionable from a physics point of view, since the entire deformation and force field should be the organizer of development, and one can hardly ascribe such a role to a single small area of the embryo body. In the same spirit, the concept of positional information encapsulated in concentration of chemicals seems questionable since the deformation and force fields in embryonic tissues are tensors. Finally, the concept of a development organized in space along three orthogonal (Cartesian) axes associated to chemical gradients seems also questionable, since early embryo development is driven by complex vortex fields, with hyperbolic trajectories which span the entire embryo. Such hyperbolic trajectories are best understood by a description in terms of dipolar components of the morphogenetic forces, whose projections along orthogonal axe have no specific meaning except as a mathematical tool. I review here the present state of description of several aspects of tetrapods morphogenesis and evolution, from the point of view of physics. It is getting clear that several basic features of tetrapods body are a direct consequences of fundamental laws of physics. Several lines of work reviewed here show that the topology of the tetrapods may be directly related to the structure of the earliest movements in embryos. The bio-mechanical approach leads to important consequences for the constraints on evolution of the craniates. Such consequences have received a controversial welcome in the last decade, although they may encapsulate the true origin of craniates, esp. simians, and eventually homo.