Intersegmental Coordination of Central Neural Oscillators for Rhythmic Movements of the Walking Legs in Crayfish, Pacifastacus Leniusculus

Abstract

ABSTRACT The neuronal circuits underlying rhythmical movements of the walking legs in crayfish Pacifastacus leniusculus are organized into central oscillators within each thoracic hemiganglion. These segmental networks are coupled in different modes of coordination during activities such as walking and limb waving. This paper examines the nature of the coupling between hemisegmental oscillators in the absence of sensory feedback, and the effect of phasic input from a major limb proprioceptor. In isolated preparations of the thoracic nerve cord, adjacent ipsilateral hemiganglia can generate a rhythmic motor output pattern in which homologous motor roots discharge approximately synchronously (the ‘in-phase’ rhythm). In contrast, contralateral hemiganglia show no evidence of any such phase-locking in their activity. A single proprioceptor at the base of each limb, the thoracic-coxal muscle receptor organ (TCMRO), can influence the timing and intensity of the rhythmic output of two or more ipsilateral thoracic ganglia. Rhythmical stretch-release of the TCMRO of the fourth ganglion on one side can entrain the motor rhythms of both the third and fourth ipsilateral hemiganglia, in the in-phase pattern of coordination. The TCMRO also elicits intra- and intersegmental reflexes during expression of the in-phase motor rhythm. These reflexes are centrally modulated in a phasedependent manner. Thus mechanical stimulation of the TCMRO co-activates homologous sets of motoneurones in adjacent ipsilateral hemiganglia only during their active (spiking) phase in the motor rhythm. We discuss the implications of these results for the generation and coordination of walking and other rhythmic behaviour of the thoracic limbs in decapod Crustacea. Our data shed further light on the properties of the motor rhythm generator for walking in crayfish, and suggest that the concept of a central pattern generator cannot be rigorously applied in this system.