Pattern formation during insect leg segmentation: Studies with a prepattern of a cell surface antigen

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Abstract
A monoclonal antibody (MAb) that binds to a cell surface antigen selectively localized to epithelial cells undergoing morphogenesis was used to study the segmentation of the growing embryonic leg of the cockroachPeriplaneta americana. The MAb labels circumferential stripes of cells at locations where invagination will occur to form the leg segments. The formation of these stripes precedes any morphological change in the epithelial layer or in individual cells. The temporal and spatial distribution of the antigen indicates the existence of a prepattern for leg segmentation, examination of which can give information about pattern generating mechanisms. Although highly stereotyped, the sequence in which the stripes appear does not follow a simple pattern proceeding in one direction along the proximal-distal axis. It is proposed that each stripe is a boundary in a positional field. Stripe formation leads to the division of the leg into a repeating series of identical positional fields. Three different mechanisms for the formation of stripes of MAb labeled cells have been observed and the role of each in the evolution of the insect leg is discussed. Measurements of leg and leg segment lengths when the various stripes appear has demonstrated considerable variation, particularly at the early stages of segmentation. Rules or mechanisms generating pattern at early stages of development are not rigid. Variations arising are compensated for by later occurring events so that stereotyped structures are formed.