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R Kittmann

Publications and source records attributed to R Kittmann.

6 recordsLinked to original sources

The antennal motor system of the stick insect Carausius morosus: anatomy and antennal movement pattern during walking.

The stick insect Carausius morosus continuously moves its antennae during locomotion. Active antennal movements may reflect employment of antennae as tactile probes. Therefore, this study treats two basic aspects of the antennal motor system: First, the anatomy of antennal joints, muscles, nerves and motoneurons is described and discussed in comparison with other species. Second, the typical movement pattern of the antennae is analysed, and its spatio-temporal coordination with leg movements described. Each antenna is moved by two single-axis hinge joints. The proximal head-scape joint is controlled by two levator muscles and a three-partite depressor muscle. The distal scape-pedicel joint is controlled by an antagonistic abductor/ adductor pair. Three nerves innervate the antennal musculature, containing axons of 14-17 motoneurons, including one common inhibitor. During walking, the pattern of antennal movement is rhythmic and spatiotemporally coupled with leg movements. The antennal abduction/adduction cycle leads the protraction/retraction cycle of the ipsilateral front leg with a stable phase shift. During one abduction/adduction cycle there are typically two levation/depression cycles, however, with less strict temporal coupling than the horizontal component. Predictions of antennal contacts with square obstacles to occur before leg contacts match behavioural performance, indicating a potential role of active antennal movements in obstacle detection.

Animals↗

The femur-tibia control system in a proscopiid (Caelifera, Orthoptera): a test for assumptions on the functional basis and evolution of twig mimesis in stick insects.

The extremely slow return movements observed in stick insects (phasmids) after imposed changes in posture are termed catalepsy. In the literature, catalepsy is treated as a behavioural component of the twig mimesis observed in walking stick insects. It is produced by the high gain of the velocity-sensitive component of the relevant joint control systems and by the non-linear dependency of its time constant on movement velocity. The high gain, in turn, causes the system to work close to instability, and this may have driven the evolution of gain control mechanisms. Although these statements represent plausible assumptions, based on correlated occurrence, they remain largely hypothetical like many ideas concerning evolutionary tendencies. To test these hypotheses, we studied catalepsy and the relevant properties of the femur-tibia control system in the middle and hind legs of Prosarthria teretrirostris.cf. Prosarthria teretrirostris is a proscopiid closely related to grasshoppers and locusts. With its slender, green-to-brown body and legs, it shows clear morphological twig mimesis, which has evolved independently of the well-known twig mimesis in stick insects. The animals show clear catalepsy. The main properties of femur-tibia joint control are remarkably similar between proscopiids and stick insects (e.g. the marked sensitivity to movement velocity rather than to joint position and the non-linear dependency of the time constants of response decay on movement velocity), but there are also important differences (habituation and activity-related mechanisms of gain control are absent). Together, these results validate the main concepts that have been developed concerning the neural basis and evolution of catalepsy in stick insects and its relationship to twig mimesis, while demonstrating that ideas on the role of habituation and gain control should be refined.

Animals↗

Neural mechanisms of adaptive gain control in a joint control loop: muscle force and motoneuronal activity

An adaptive gain control system of a proprioceptive feedback system, the femur­tibia control loop, is investigated. It enables the joint control loop to work with a high gain but it prevents instability oscillations. In the inactive stick insect, the realisation of specific changes in gain is described for tibial torque, for extensor tibiae muscle force and for motoneuronal activity. In open-loop experiments, sinusoidal stimuli are applied to the femoral chordotonal organ (fCO). Changes in gain that depend on fCO stimulus parameters (such as amplitude, frequency and repetition rate), are investigated. Furthermore, spontaneous and touch-induced changes in gain that resemble the behavioural state of the animal are described. Changes in gain in motoneurones are always realised as changes in the amplitude of modulation of their discharge frequency. Nevertheless, depending on the stimulus situation, two different mechanisms underlie gain changes in motoneurones. (i) Changes in gain can be based on changes in the strength of the sensorimotor pathways that transmit stimulus-modulated information from the fCO to the motoneurones. (ii) Changes in gain can be based on changes in the mean activity of a motoneurone by means of its spike threshold: when, during the modulation, the discharge of a motoneurone is inhibited for part of the stimulus cycle, then a change in mean activity subsequently causes a change in modulation amplitude and gain. A new neuronal mechanism is described that helps to compensate the low-pass filter characteristics of the muscles by an increased activation, especially by a sharper distribution of spikes in the stimulus cycle at high fCO stimulus frequencies.

Journal Article↗

Premotor interneurons in generation of adaptive leg reflexes and voluntary movements in stick insects.

We investigated the role of local nonspiking interneurons involved in motor control of legs in the stick insects, Carausius morosus. In a preparation that allowed the animals to perform active leg movements such as adaptive tactile reflexes, proprioceptive reflexes, and walking, we gathered the following results. Almost all tested nonspiking interneurons that provide synaptic drive onto moto-neurons of the proximal leg muscles contribute to all of the motor programs underlying tactile reflexes and voluntary leg movements such as walking, searching, and rocking. Most of them are also involved in the generation of proprioceptive reflexes. All motor programs for coactivation, avoidance reflexes, resistance reflexes, and voluntary leg movements result from parallel pathways including nonspiking interneurons that support and others that oppose the motoneuronal activity. The contribution of a single interneuron to the different motor program is specific: it can be supporting for one motor program but opposing for the other. Even for the same motor program, for example, coactivation, the contribution of an individual interneuron can depend on the stimulus site from where the response is elicited. Our results support the idea that the different motor patterns for adaptive tactile reflexes, resistance reflexes, and voluntary leg movements emerge from a multifunctional neuronal circuit that is reorganized corresponding to the motor behavior performed. The actual motor pattern is then shaped by distributed information processing in parallel supporting and opposing pathways.

Acclimatization↗

Octopamine effects mimick state-dependent changes in a proprioceptive feedback system.

The modulatory actions of the biogenic amine octopamine on the femur tibia (FT) control loop in the stick insect Carausius morosus were examined. The response properties of the FT control loop were determined under open loop conditions. Mechanical stimulation of the femoral chordotonal organ (fCO) was the input and tibial movement and motoneuronal activity were measured as the output of the system. Following octopamine injection into the hemolymph of intact, inactive animals, two consecutive phases occurred at the behavioral level. Octopamine caused initially an activation of the animal. During this first phase (3.5-12 min duration) the response properties of the FT control loop were similar to those found in animals that were activated by tactile stimuli under normal conditions. Afterward, animals became inactive. During this second phase (15-20 min duration), the gain of the control loop was zero and no resistance reflex in the FT joint was generated in response to fCO stimulation. However, active movements of the tibia could still be elicited. As we could show in restrained animals, where DL-octopamine was applied topically onto the undesheated mesothoracic ganglion, the complete suppression of the resistance reflex on the motoneuronal level was dose dependent starting at concentrations of 5 x 10(-3) M octopamine. We could show that octopamine specifically suppressed the pathways involved in the resistance reflex, while feedback loop responses to fCO stimuli typical for active animals could still be elicited. Our results indicate that an increase in the octopamine concentration mimicks activation of the animal: Properties being characteristic for the control of the FT joint in the inactive animal are inhibited by octopamine, while properties of the FT control loop typical for the active animal appear to be facilitated following octopamine injection. The results clearly demonstrate that different pathways in the neuronal network underlying the FT control loop are involved in the responses of the control loop to fCO stimuli in the inactive and active behavioral states of the stick insect.

Animals↗

Intracellular recordings from nonspiking interneurons in a semiintact, tethered walking insect.

Nonspiking interneurons were investigated in a tethered, walking insect, Carausius morosus, that was able to freely perform walking movements. Experiments were carried out with animals walking on a lightweight, double-wheel treadmill. Although the animal was opened dorsally, the walking system was left intact. Intracellular recordings were obtained from the dorsal posterior neuropil of the mesothoracic ganglion. Nonspiking interneurons, in which modulations of the membrane potential were correlated with the walking rhythm, were described physiologically and stained with Lucifer Yellow. Interneurons are demonstrated in which membrane potential oscillations mirror the leg position or show correlation with the motoneuronal activity of the protractor and retractor coxae muscles during walking. Other interneurons showed distinct hyperpolarizations at certain important trigger points in the step cycle, for example, at the extreme posterior position. Through electrical stimulation of single, nonspiking interneurons during walking, the motoneuronal activity in two antagonistic muscles--protractor and retractor coxae--could be reversed and even the movement of the ipsilateral leg could be influenced. The nonspiking interneurons described appear to be important premotor elements involved in walking. They receive, integrate, and process information from different leg proprioceptors and drive groups of leg motoneurons during walking.

Animals↗