Variability of blowfly head optomotor responses

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Abstract
SUMMARY: Behavioural responses of an animal are variable even when the animal experiences the same sensory input several times. This variability can arise from stochastic processes inherent to the nervous system. Also, the internal state of an animal may influence a particular behavioural response. In the present study, we analyse the variability of visually induced head pitch responses of tethered blowflies by high-speed cinematography. We found these optomotor responses to be highly variable in amplitude. Most of the variability can be attributed to two different internal states of the flies with high and low optomotor gain, respectively. Even within a given activity state, there is some variability of head optomotor responses. The amount of this variability differs for the two optomotor gain states. Moreover, these two activity states can be distinguished on a fine timescale and without visual stimulation, on the basis of the occurrence of peculiar head jitter movements. Head jitter goes along with high gain optomotor responses and haltere oscillations. Halteres are evolutionary transformed hindwings that oscillate when blowflies walk or fly. Their main function is to serve as equilibrium organs by detecting Coriolis forces and to mediate gaze stabilisation. However, their basic oscillating activity was also suggested to provide a gain-modulating signal. Our experiments demonstrate that halteres are not necessary for high gain head pitch to occur. Nevertheless, we find the halteres to be responsible for one component of head jitter movements. This component may be the inevitable consequence of their function as equilibrium and gaze-stabilising organs.