The Locust Tegula: Significance for Flight Rhythm Generation, Wing Movement Control and Aerodynamic Force Production

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Abstract
The tegula, a complex sense organ associated with the wing base of the locust, plays an important role in the generation of the flight motor pattern. Here its function in the control of wing movement and aerodynamic force production is described. The vertical component of forewing movement was monitored while recording intracellularly from flight motoneurones during stationary flight. First, in accordance with previous electrophysiological results, stimulation of hindwing tegula afferents was found to reset the wingstroke to the elevation phase in a well-coordinated manner. Second, recordings made before and after removal of fore- and hindwing tegulae were compared. This comparison demonstrated that the delayed onset of elevator motoneurone activity caused by tegula removal is accompanied by a corresponding delay in the upstroke movement of the wings. The consequences of this delayed upstroke for aerodynamic force production were investigated by monitoring wing movements and lift generation simultaneously. A marked decrease in net lift generation was observed following tegula removal. Recordings of wing pronation indicate that this decrease in lift is primarily due to the delayed upstroke movement - that is, to a delay of the wings near the aerodynamically unfavourable downstroke position. It is concluded that the tegula of the locust hindwing signals to the nervous system the impending completion of the wing downstroke and allows initiation of the upstroke movement immediately after the wings have reached the lower reversal point of the wingstroke. The functional significance of tegula feedback and central rhythm generation for locust flight control are discussed.