The control of wing kinematics and flight forces in fruit flies (Drosophila spp.).

By simultaneously measuring flight forces and stroke kinematics in several species of fruit flies in the genus Drosophila, we have investigated the relationship between wing motion and aerodynamic force production. We induced tethered flies to vary their production of total flight force by presenting them with a vertically oscillating visual background within a closed-loop flight arena. In response to the visual motion, flies modulated their flight force by changing the translational velocity of their wings, which they accomplished via changes in both stroke amplitude and stroke frequency. Changes in wing velocity could not, however, account for all the modulation in flight force, indicating that the mean force coefficient of the wings also increases with increasing force production. The mean force coefficients were always greater than those expected under steady-state conditions under a variety of assumptions, verifying that force production in Drosophila spp. must involve non-steady-state mechanisms. The subtle changes in kinematics and force production within individual flight sequences demonstrate that flies possess a flexible control system for flight maneuvers in which they can independently control the stroke amplitude, stroke frequency and force coefficient of their wings. By studying four different-sized species, we examined the effects of absolute body size on the production and control of aerodynamic forces. With decreasing body size, the mean angular wing velocity that is required to support the body weight increases. This change is due almost entirely to an increase in stroke frequency, whereas mean stroke amplitude was similar in all four species. Despite the elevated stroke frequency and angular wing velocity, the translational velocity of the wings in small flies decreases with the reduction in absolute wing length. To compensate for their small size, D. nikananu must use higher mean force coefficients than their larger relatives.