The development of sensory nerves within the wing ofDrosophila melanogaster : A light- and electron-microscope study.

TheDrosophila wing is a simple structure bearing many sensory neurons which form an elementary pattern of nerves within the veins, and provides a simple system for studying the formation of nerve pathways. A light-and electron-microscope study of the developing pupal wing was undertaken to establish the time at which the different classes of sensillum differentiate and to determine the arrangement of tissues within the wing during sensory axon out-growth in order to assess possible candidates for directing axon outgrowth. Major findings were 1. Wing development passes through three main stages: at 6-12 h after puparium formation the wing is flantened and secreting pupal cuticle; at 15-18 h the wing is dramatically inflated but is still a simple epithelium secreting pupal cuticle; at 21-24 h the wing collapses and begins to differentiate adult structures. 2. There are no persisting larval nerves which might act as pioneers or pathfinders for the later developing adult neurons. 3. Axon bundles are first observed during the 15-18 h stage prior to the pupal moult when the wing is still secreting pupal cuticle. 4. At this stage the wing is an inflated sac, without any veins or orderly arrangements of tracheae which might act as guides for axon outgrowth. Vein formation takes placeafter formation of the axon pathways. 5. The bristle axons grow along the anterior wing margin in close contact with the basal lamina of the epithelial cells, often within a gap between the processes of the epithelial cells, which could mechanically channel their out-growth. 6. The campaniform sensillum axons appear to navigate along the inner surface of the wing epithelium rather like the pioneer axons found in embryonic appendages of other insects. 7. Differentiation of sensory neurons takes place long before differentiation of the other cells associated with sensilla; most axons are present by 18 h when the wing is still secreting pupal cuticle but morphological differentiation of the bristle shaft and socket cells is not observed until about 42 h when the wing is secreting the cuticulin layer of the adult cuticle.