Mapping motor neurone activity to overt behaviour in the leech: internal pressures produced during locomotion

Several behaviour patterns have been studied in the leech at both the kinematic and neuronal levels. However, very little is known about how patterns of motor neurone activity map to actual movements. Internal pressure is an essential biomechanical property in this process, being responsible for producing the rigidity and posture that allow the directed delivery of forces produced by muscle contraction. To obtain a better understanding of the biomechanical processes involved in movement of the leech, we have measured the internal pressure of the animal by placing catheters through the body wall and into the gut of intact animals showing normal patterns of behaviour. Each type of behaviour had a characteristic pressure waveform. The elongation phase of crawling produced a rapid increase in pressure that peaked when midbody segments were maximally elongated. The pressure produced during the contraction phase of crawling depended on the type of crawl, only inchworm crawling producing a second peak. Whole-body shortening in response to a head poke also produced a pressure peak, but it had a faster rise time. Swimming produced the largest pressure, which was marked by a large sustained increase that fluctuated phasically with undulations of the body. Dual pressure recordings using two catheters demonstrated that pressure was not uniform along the length of the leech, indicating that the body cavity is functionally compartmentalised. Injecting fluid into the gut via a recording catheter allowed us to determine the effects of increasing internal volume on pressure. In line with previous predictions made using an abstract biomechanical model of the leech hydroskeleton, we found that an increase in the volume caused a reduction in the pressure. We are in the process of constructing a more realistic biomechanical model of the leech, based on actual data reported elsewhere. The results in this paper will provide key tests for refining these models.