Cell sociology and the problem of position effect: Pattern formation, origin and role of gradients

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Abstract
The control of pattern formation and the significance of gradients is reconsidered on the basis of the concept of cell sociology (which takes into account continuous exchange of information between cells and the possibility of autonomous progression in differentiation). Not all traits of a pattern are imposed by a single prepattern, which would be an organized molecular framework or a gradient. Patterns are unfolded in steps; these are readjustments of a cell population to intrinsic and extrinsic changes in cell activities. Prepatterns are the various components of the programme of every readjustment and are established by information of various origins, which can be dissociated experimentally: determination (elementary social prepattern), pre-existing organization (antecedent pp.), surrounding cell populations (environmental pp.), position among other tissues (positional pp.) and the organization of inducers (imprinting pp.). Every transitory pattern formed during a readjustment serves as antecedent pp. during the next readjustment. Covert graded patterns result from various aspects of the social behaviour of cells (growth, aggregation, induction, cell renewal) and may serve as antecedent or imprinting prepatterns. They appear as ‘water marks’ in the final patterns, or generate overt graded patterns. They also manifest themselves in temporal patterns, particularly in gradients of relative growth.