Adaptive plasticity of spino-extraocular motor coupling during locomotion in metamorphosing Xenopus laevis.

During swimming in the amphibian ITALIC! Xenopus laevis, efference copies of rhythmic locomotor commands produced by the spinal central pattern generator (CPG) can drive extraocular motor output appropriate for producing image-stabilizing eye movements to offset the disruptive effects of self-motion. During metamorphosis, ITALIC! X. laevisremodels its locomotor strategy from larval tail-based undulatory movements to bilaterally synchronous hindlimb kicking in the adult.

Spinal efference copy signaling and gaze stabilization during locomotion in juvenile Xenopus frogs.

In swimming Xenopus laevis tadpoles, gaze stabilization is achieved by efference copies of spinal locomotory CPG output that produce rhythmic extraocular motor activity appropriate for minimizing motion-derived visual disturbances. During metamorphosis, Xenopus switches its locomotory mechanism from larval tail-based undulatory movements to bilaterally synchronous hindlimb kick propulsion in the adult. The change in locomotory mode leads to body motion dynamics that no longer require conjugate left-right eye rotations for effective retinal image stabilization.

Premotor interneurons in the local control of stepping motor output for the stick insect single middle leg.

In insect walking systems, nonspiking interneurons (NSIs) play an important role in the control of posture and movement. As such NSIs are known to contribute to state-dependent modifications in processing of proprioceptive signals from the legs. For example, NSIs process a flexion of the femur-tibia (FTi) joint signaled by the femoral chordotonal organ (fCO) such that the stance phase motor output is reinforced in the active locomotor system.

Control of stepping velocity in the stick insect Carausius morosus.

We performed electrophysiological and behavioral experiments in single-leg preparations and intact animals of the stick insect Carausius morosus to understand mechanisms underlying the control of walking speed. At the level of the single leg, we found no significant correlation between stepping velocity and spike frequency of motor neurons (MNs) other than the previously shown modification in flexor (stance) MN activity. However, pauses between stance and swing motoneuron activity at the transition from stance to swing phase and stepping velocity are correlated.

Slow temporal filtering may largely explain the transformation of stick insect (Carausius morosus) extensor motor neuron activity into muscle movement.

Understanding how nervous systems generate behavior requires understanding how muscles transform neural input into movement. The stick insect extensor tibiae muscle is an excellent system in which to study this issue because extensor motor neuron activity is highly variable during single leg walking and extensor muscles driven with this activity produce highly variable movements. We showed earlier that spike number, not frequency, codes for extensor amplitude during contraction rises, which implies the muscle acts as a slow filter on the time scale of burst interspike intervals (5-10 ms).

Different motor neuron spike patterns produce contractions with very similar rises in graded slow muscles.

Graded muscles produce small twitches in response to individual motor neuron spikes. During the early part of their contractions, contraction amplitude in many such muscles depends primarily on the number of spikes the muscle has received, not the frequency or pattern with which they were delivered. Stick insect (Carausius morosus) extensor muscles are graded and thus would likely show spike-number dependency early in their contractions.

Natural neural output that produces highly variable locomotory movements.

We recorded fast extensor tibiae motor neuron activity during single-legged treadmill walking in the stick insect, Carausius morosus. We used this activity to stimulate the extensor muscle motor nerve, observed the resulting extensor muscle contractions under isotonic conditions, and quantified these contractions with a variety of measures. Extensor contractions induced in this manner were highly variable, with contraction measures having SDs of 12 to 51%, and ranges of 82 to 275%, when expressed as percentages of the means, an unexpectedly wide range for a locomotory pattern.