Somitogenesis in amphibia
A somite pre-pattern is established shortly before visible segmentation. The pre-pattern results from the interaction of two components: a wave of cell behavioural change that passes along the axis, and, an underlying co-ordination of the cells that is the basis for their association into large somite-sized groupings. The evidence is derived from studies of the zones of abnormal segmentation that follow temperature shocks delivered between the neurula and tail-bud stages (Pearson & Elsdale, 1979). Temperature shock given earlier at the mid-gastrula stage is however ineffective in inducingabnormalities in somitogenesis. Shocks given before the mid-gastrula stage reveala prior period of sensitivity stretching back into the blastula. Thus early and late sensitive periods can be defined separated by a short refactory period. Quite different patterns in the distribution of somite abnormalities characterize the results of shock during the two sensitive periods, suggesting different aetiologies. It is concluded that the wave of rapid cell change is set up early in embryogenesis during theblastula stage, and each cell of the prospective paraxial mesoderm carries a determination to change after a specific length of time, i.e. a countdown is set in each cell. As a result of the movements of gastrulation, the prospective paraxial mesoderm cells become laid out along the axis of the neurula in the order (antero-posterior sequence) in which they will change. The achievement of the correct redistribution of the cells depends crucially on the conservation of the sequence in the blastula by the maintenanceof topological integrity throughout gastrulation. It is suggested that early shock disturbs gastrulation movements, causing some mixing up of the cells resulting in incoherenceof the wavefront. Whereas early shocks are thus assumed to affect the wave, the evidencesuggests that late shock undoes co-ordination. It is concluded therefore that co-ordination is established later, after the refractory period, around the late gastrula stage.