Models for positional signalling with application to the dorsoventral patterning of insects and segregation into different cell types.

Models of pattern formation and possible molecular realizations are discussed and compared with recent experimental observations. In application to the dorsoventral patterning of insects, it is shown that a superposition of two pattern-forming reactions is required. The first system generates the overall dorsoventral polarity of the oocyte, the second generates the positional information proper with a stripe-like region of high concentration along the ventral side of the embryo. A single reaction would be insufficient since the two reactions require different parameters. The model accounts for the orientation of the DV axes of the oocytes in the ovary of Musca domestica and Sarcophaga, independent of the DV axis of the mother, for the formation of several ventral furrows in the absence of the primary gurken/torpedo system in Drosophila, as well as for the good size regulation of the dorsoventral axis as observed in some insect species. Segregation of a homogeneous cell population into different cell types requires autocatalytic processes that saturate at relatively low concentrations and nondiffusible substances responsible for the autocatalytic feed-back loops. Thus, these loops can be realized directly on the gene level via their gene products, for instance, by the mutual repression of two genes. A balance of the two cell types is achieved by a long-ranging substance interfering with the self-enhancing process. This substance is expected to have a more or less homogeneous distribution. This model accounts for the reestablishment of the correct proportion after an experimental interference and the change of determination after transplantation. Applications to the segregation of pre-stalk and prespore cells in Dictyostelium and of neuroblast cells from the ventral ectoderm in Drosophila are provided.