Epidermal feet in pupal segment morphogenesis

Epidermal cells in insect integumental epithelia develop branched cytoskeletal extensions or feet at their base that are similar in appearance to the processes put out by cells in tissue culture. We have developed a procedure to show the feet that gives an effect as if thousands of cells randomly arranged in the epithelium had each been injected with a lead salt visualized as black lead sulphide. The procedure depends upon the fact that after brief glutaraldehyde fixation, tannic acid only penetrates some cells where it mordants lead ions and binds osmium. Individual cells visualized in this manner show their outlines as if they are separate in a tissue culture although they are part of a closely packed epithelium. The feet are metamorphic structures formed after pupal commitment and are necessary for metamorphic changes in segment shape. In Calpodes larvae the feet are orientated axially in the direction of the segmentally repeating gradient and may extend for several cell diameters. They extend under the influence of low titres of 20-hydroxyecdysone such as those occurring in the intermoult. When stimulated by high titres like those in pre-pupae, the feet contract at the same time as the segments shorten to pupal proportions. We believe that cell processes like the epidermal feet are ubiquitous but that they have often been overlooked because of the difficulty of demonstrating the outlines of single cells that are united in epithelia.