The development and evolution of crossveins in insect wings.

The formation of crossveins in Drosophila was an important early case study in understanding the role of the environment in the development and evolution of morphological structures. More recent work has shown that signalling processes play a crucial role in the formation of crossveins in Drosophila and that the interaction of a heat shock factor, Hsp90, with components of signal transduction pathways may account for the sensitivity of these structures to environmental perturbations. A new model for the development of crossveins is presented that divides the formation of crossveins into 3 separate stages. First, the number and placement of the crossveins is determined by signalling along the proximal-distal axis of the wing. This signal may involve the cdc42 gene product and the Jun-N-terminal Kinase signal transduction pathway. Then, during the second stage, an inductive signal from the dorsal wing epithelium is sent to the ventral wing epithelium at locations specified by the first signal. The second signal appears to involve the BMP-like signalling pathway. Finally, in the third stage, a domain of vein competent cells is defined by the signalling from the EGF-receptor-Map Kinase signal transduction pathway, and the exact location of the veins is eventually determined within that domain by Notch-Delta signalling interactions. By altering components of these 3 stages, insects can independently regulate the presence or absence, the number and placement, and the thickness and flexibility of the crossveins. This capacity for the modulation of crossvein structure in many different ways may have contributed to the evolution of different modes of insect flight.