Active tactile exploration and tactually induced turning in tethered walking stick insects.

Many animals use their tactile sense for active exploration and tactually guided behaviors such as near-range orientation. In insects, tactile sensing is often intimately linked to locomotion, resulting in the orchestration of several concurrent active movements, including turning of the entire body, rotation of the head, and searching or sampling movements of the antennae. The present study aims at linking the sequence of tactile contact events to associated changes of all three kinds of these active movements (body, head and antennae). To do so, we chose the Indian stick insect Carausius morosus, an organism commonly used to study sensory control of locomotion. Methodologically, we combined recordings of walking speed, heading, whole-body kinematics and antennal contact sequences during stationary, tethered walking and controlled presentation of an 'artificial twig' for tactile exploration. Our results show that object presentation episodes as brief as 5 s are sufficient to allow for a systematic investigation of tactually induced turning behavior in walking stick insects. Animals began antennating the artificial twig within 0.5 s, and altered the beating fields of both antennae in a position-dependent manner. This change was mainly carried by a systematic shift of the head-scape joint movement and accompanied by associated changes in contact likelihood, contact location and sampling direction of the antennae. The turning tendency of the insect also depended on stimulus position, whereas the active, rhythmic head rotation remained unaffected by stimulus presentation. We conclude that the azimuth of contact location is a key parameter of active tactile exploration and tactually induced turning in stick insects.

Speed-dependent interplay between local pattern-generating activity and sensory signals during walking in Drosophila.

In insects, the coordinated motor output required for walking is based on the interaction between local pattern-generating networks providing basic rhythmicity and leg sensory signals, which modulate this output on a cycle-to-cycle basis. How this interplay changes speed-dependently and thereby gives rise to the different coordination patterns observed at different speeds is not sufficiently understood. Here, we used amputation to reduce sensory signals in single legs and decouple them mechanically during walking in Drosophila. This allowed for the dissociation between locally generated motor output in the stump and coordinating influences from intact legs. Leg stumps were still rhythmically active during walking. Although the oscillatory frequency in intact legs was dependent on walking speed, stumps showed a high and relatively constant oscillation frequency at all walking speeds. At low walking speeds we found no strict cycle-to-cycle coupling between stumps and intact legs. In contrast, at high walking speeds stump oscillations were strongly coupled to the movement of intact legs on a one-to-one basis. Although during slow walking there was no preferred phase between stumps and intact legs, we nevertheless found a preferred time interval between touch-down or lift-off events in intact legs and levation or depression of stumps. Based on these findings, we hypothesize that, as in other insects, walking speed in Drosophila is predominantly controlled by indirect mechanisms and that direct modulation of basic pattern-generating circuits plays a subsidiary role. Furthermore, inter-leg coordination strength seems to be speed-dependent and greater coordination is evident at higher walking speeds.

A laser-supported lowerable surface setup to study the role of ground contact during stepping.

We introduce a laser-supported setup to study the influence of afferent input on muscle activation during walking, using a movable ground platform. This approach allows investigating if and how the activity of stance phase muscles of an insect (e.g. stick insect) responds to a missing ground contact signal. The walking surface consists of a fixed and a lowerable part, which can be lowered to defined levels below the previous ground level at any time point during a walking sequence. As a consequence, the leg under investigation finds either a lower ground level or no ground support at all. The lowerable walking surface consists of a 49 mm × 34 mm stainless steel surface, made slippery and equipped for tarsal contact monitoring, similar to the system that was described by Gruhn and colleagues (Gruhn et al., 2006). The setup controller allows pneumatic lowering of the surface and subsequent detection of tarsal entry into the previous ground level with the help of a thin sheet of laser light and a corresponding detector. Here, we describe basic properties of the new setup and show the results of first experiments to demonstrate its use for the study of sensory and central influences in stepping of a small animal. In the experiments, we compare the effect of ground-support ("control") with either steps into the hole (SiH), ground support at a lower surface level, or the amputation of the tarsus on the onset of EMG activity in the flexor tibiae muscle of the stick insect.