Pattern and polarity of sclerites in adult abdominal segments of Calliphora erythrocephala (Diptera): anlage rotation experiments

The anlagen of imaginal histoblasts in the abdominal segments of Calliphora (higher Diptera) present an interesting problem, which bears on recent concepts employed in the consideration of spatial patterning in insects. They differ from imaginal discs with respect to larval organization and activity, and in the absence of the progressive pattern of genetic determination during the larval period, characteristic of imaginal discs. How is the adult pattern in the abdominal segments determined? The experiments presented here seek to clarify the spatial parameters involved in control of adult pattern and polarity in the adult segment. A series of 180° rotations of hypodermal grafts bearing the anlagen singly, or in combination, or of larval intersegmental hypodermis, indicate that polarity is determined within the anlagen, through interaction with local larval epidermis either before or during histoblast migration. The nature of the sclerites, too, is primarily carried by the anlagen rather than determined by intersegmental information. The whole question of ‘determination of polarity’ is set out more carefully than hitherto in the light of (a) observations of the movement of epidermal cells in other systems in response to disturbance of pattern, and (b) the obvious vectorial nature of the phenomenon, which cannot be a genetic matter, but one of cell axes and of the relation of cells to segment/organism. The demonstration that (i) hemitergite and hemisternite are primarily determined by the anlagen themselves, and not by larval intersegmental membranes; and (ii) evidence indicates an influence of epidermal cells of the larva on the differentiation (as well as polarity) of imaginal histoblasts, leads to the conclusion that neither of two models considered will account for the establishment of the adult abdominal pattern among the histoblasts at metamorphosis. These models are (a) of a segmental gradient, set by the intersegmental boundaries of the previous instar, to which imaginal cells respond by interpretation of positional information; and (b) of progressive compartmentalization of pattern within the anlagen, without reference to epidermal context.