Nucleolar cycles during the fifth stadium in Manduca epidermis

The changing pattern of nucleolar structure in the epidermal cells of Manduca sexta has been correlated with hormonal changes taking place during the fifth stadium. The epidermal nucleoli show three cycles of development, the first and third of which occur at the beginnings of the intermoult and moult phases respectively and are related to larval and pupal syntheses. The second phase occurs in the middle of the stadium but prior to the onset of wandering and commitment to pupation. A phase of mitosis separates the second and third cycles. The three cycles thus correspond in time to those found in Calpodes. The three cycles of nucleolar change are superimposed over nuclear changes relating to the degree of ploidy. Each phase begins with an expansion of the condensed nucleoli to form lobed rings and then necklaces. In the first phase (day 0-3), the rings and necklaces progress to form threaded networks. Both rings and networks have many ribosomal precursor granules that are lacking in condensed nucleoli. The rings and networks are therefore presumed to be more active in rRNA synthesis than the condensed state. The first and third phases of nucleolar change occur after elevated titres of haemolymph ecdysteroid. Post-thoracic ligation of animals at ecdysis blocks nucleolar changes as well as the appearance of polyploid nuclei. Nucleolar changes may be a primary response of the epidermis to stimulation by ecdysone.