Resistance reflex that maintains upright head posture in the flesh fly neobellieria bullata (Sarcophagidae)

In flesh flies Neobellieria bullata, we investigated a resistance reflex that maintains upright head posture around the roll axis relative to the thorax. The gain of the reflex depends upon the fly's behavioral state: moving flies immediately correct 90 % of the amplitude of experimentally imposed roll perturbations, returning the head almost to the fully upright position; motionless flies allow perturbations to persist for minutes before correcting only 70 % of perturbation amplitude. To investigate the role of various neural pathways, we examined the control of head posture after sectioning relevant propriosensory or motor nerves. Excision of the prosternal chordotonal organ causes no decrements in the control of head posture. Unilateral deafferentation of a cervical propriosensory organ, the prosternal organ, induces roll towards the cut side. Unilateral section of the frontal nerve, a mixed motor nerve that supplies the neck depressors and levators, leads to unilateral deficits in correcting perturbations towards the contralateral side. After bilateral propriosensory or frontal motor nerve section, approximately 40 % of perturbation amplitude is still corrected. To determine the contributions of the passive elastic properties of the neck skeleto-muscular system, flies were tested under reversible nitrogen anesthesia. They immediately corrected 40 % of perturbation amplitude. Taken together, the results demonstrate that passive elasticity plus active prosternal nerve afference to contralateral depressors innervated by the frontal nerve in combination constitute a sufficient and necessary reflex loop to control head roll posture.