Contralateral Coordination and Retargeting of Limb Movements During Scratching in the Locust

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ABSTRACT Locusts, Schistocerca gregaria, in common with many limbed vertebrates, can make directed scratching movements in response to tactile stimulation. For instance, stimulation of different sites on a wing elicits different movements that are accurately targeted so that the hindleg tarsus passes across the stimulus site. I have analysed these limb movements to define the ability of a locust to target stimulus sites correctly under a range of experimental conditions. In particular, I describe aspects of the behaviour that reveal possible neuronal pathways underlying the responses. These neuronal pathways will be the subject of further physiological analyses. Limb targeting during scratching is continuously graded in form; different patterns of movement are not separated by sharp transitions. The computation of limb trajectory takes into account the starting posture of the hindleg, so that different trajectories can be used to reach a common stimulus site from different starting postures. Moreover, the trajectories of the two hindlegs moving simultaneously from different starting postures in response to a single stimulus can be different, so that their tarsi converge onto the common stimulus site. Different trajectories can be used to reach a common stimulus site from the same start posture. Targeting information from a forewing is passed not only down the nerve cord to the ipsilateral hindleg but also across the nerve cord, so that the contralateral hindleg can also make directed movements. This contralateral transmission does not rely on peripheral sensory feedback. When the stimulus site moves during a rhythmical scratch, the targeting of subsequent cycles reflects this change. Both ipsilateral and contralateral hindlegs can retarget their movements. The trajectory of a single cycle of scratching directed towards a particular stimulus site can be modified after it has begun, so that the tarsus is redirected towards a new stimulus site.