scholarly journals Sensory Feedback During Active Movements of Stick Insects

1987 ◽  
Vol 133 (1) ◽  
pp. 137-156 ◽  
Author(s):  
G. WEILAND ◽  
U. T. KOCH

In the stick insect Carausius momsus, the role of the chordotonal organ was investigated using a new experimental arrangement which artificially closes the femur-tibia control system. The chordotonal organ was stimulated during voluntary movements by applying trapezoidal ramp stimuli in the closed-loop configuration. The results demonstrate that the feedback loop is used to control the end points of joint movement. In addition, it was found that the control system counteracts experimentally applied velocity changes imposed during the middle part of the movements. Acceleration-sensitive units are shown to contribute to the reaction. The results show that during active voluntary movements reflexes measured in the inactive animal are neither simply incorporated in a servo-system nor suppressed. Instead their characteristics are altered so that the voluntary movements are executed as intended by the animal. Thus reflexes cannot be considered as a fixed behavioural unit; rather their changing role must be analysed in the context of the behaviour studied.

1986 ◽  
Vol 120 (1) ◽  
pp. 369-385 ◽  
Author(s):  
G. WEILAND ◽  
U. BÄSSLER ◽  
M. BRUNNER

An experimental arrangement was constructed which is based on the open-loop femur-tibia control system of two stick insect species (Carausius morosus and Cuniculina impigra). It could be artificially closed in the following way: the position of the tibia was measured by an optical device and this value was used to drive a penmotor which moved the receptor apodeme of the femoral chordotonal organ in the same way as in intact animals. This arrangement allows direct comparison of the behaviour of the open-loop and the closed-loop system as well as introducing an additional delay. The Carausius system has a phase reserve of only 30°-50° and the factor of feedback control approaches 1 between 1 and 2 Hz. This agrees with the observation that an additional delay of 70–200 ms produces long-lasting oscillations of 1–2 Hz. The Cuniculina system has a larger phase reserve and consequently a delay of 200 ms produced no oscillations. All experiments show that extrapolation from the open-loop system to the closed-loop system is valid, despite the non-linear characteristics of the loop. Consequences for servo-mechanisms during walking and rocking movements are discussed.


2005 ◽  
Vol 93 (3) ◽  
pp. 1255-1265 ◽  
Author(s):  
Björn Ch. Ludwar ◽  
Marie L. Göritz ◽  
Joachim Schmidt

Locomotion requires the coordination of movements across body segments, which in walking animals is expressed as gaits. We studied the underlying neural mechanisms of this coordination in a semi-intact walking preparation of the stick insect Carausius morosus. During walking of a single front leg on a treadmill, leg motoneuron (MN) activity tonically increased and became rhythmically modulated in the ipsilateral deafferented and deefferented mesothoracic (middle leg) ganglion. The pattern of modulation was correlated with the front leg cycle and specific for a given MN pool, although it was not consistent with functional leg movements for all MN pools. In an isolated preparation of a pair of ganglia, where one ganglion was made rhythmically active by application of pilocarpine, we found no evidence for coupling between segmental central pattern generators (CPGs) that could account for the modulation of MN activity observed in the semi-intact walking preparation. However, a third preparation provided evidence that signals from the front leg's femoral chordotonal organ (fCO) influenced activity of ipsilateral MNs in the adjacent mesothoracic ganglion. These intersegmental signals could be partially responsible for the observed MN activity modulation during front leg walking. While afferent signals from a single walking front leg modulate the activity of MNs in the adjacent segment, additional afferent signals, local or from contralateral or posterior legs, might be necessary to produce the functional motor pattern observed in freely walking animals.


2006 ◽  
Vol 96 (6) ◽  
pp. 3532-3537 ◽  
Author(s):  
Turgay Akay ◽  
Ansgar Büschges

Reinforcement of movement is an important mechanism by which sensory feedback contributes to motor control for walking. We investigate how sensory signals from movement and load sensors interact in controlling the motor output of the stick insect femur–tibia (FT) joint. In stick insects, flexion signals from the femoral chordotonal organ (fCO) at the FT joint and load signals from the femoral campaniform sensilla (fCS) are known to individually reinforce stance-phase motor output of the FT joint by promoting flexor and inhibiting extensor motoneuron activity. We quantitatively compared the time course of inactivation in extensor tibiae motoneurons in response to selective stimulation of fCS and fCO. Stimulation of either sensor generates extensor activity in a qualitatively similar manner but with a significantly different time course and frequency of occurrence. Inactivation of extensor motoneurons arising from fCS stimulation was more reliable but more than threefold slower compared with the extensor inactivation in response to flexion signals from the fCO. In contrast, simultaneous stimulation of both sense organs produced inactivation in motoneurons with a time course typical for fCO stimulation alone, but with a frequency of occurrence characteristic for fCS stimulation. This increase in probability of occurrence was also accompanied by a delayed reactivation of the extensor motoneurons. Our results indicate for the first time that load signals from the leg affect the processing of movement-related feedback in controlling motor output.


1999 ◽  
Vol 81 (4) ◽  
pp. 1856-1865 ◽  
Author(s):  
Dietmar Hess ◽  
Ansgar Büschges

Role of proprioceptive signals from an insect femur-tibia joint in patterning motoneuronal activity of an adjacent leg joint. Interjoint reflex function of the insect leg contributes to postural control at rest or to movement control during locomotor movements. In the stick insect ( Carausius morosus), we investigated the role that sensory signals from the femoral chordotonal organ (fCO), the transducer of the femur-tibia (FT) joint, play in patterning motoneuronal activity in the adjacent coxa-trochanteral (CT) joint when the joint control networks are in the movement control mode of the active behavioral state. In the active behavioral state, sensory signals from the fCO induced transitions of activity between antagonistic motoneuron pools, i.e., the levator trochanteris and the depressor trochanteris motoneurons. As such, elongation of the fCO, signaling flexion of the FT joint, terminated depressor motoneuron activity and initiated activity in levator motoneurons. Relaxation of the fCO, signaling extension of the FT joint, induced the opposite transition by initiating depressor motoneuron activity and terminating levator motoneuron activity. This interjoint influence of sensory signals from the fCO was independent of the generation of the intrajoint reflex reversal in the FT joint, i.e., the “active reaction,” which is released by elongation signals from the fCO. The generation of these transitions in activity of trochanteral motoneurons barely depended on position or velocity signals from the fCO. This contrasts with the situation in the resting behavioral state when interjoint reflex action markedly depends on actual fCO stimulus parameters, i.e., position and velocity signals. In the active behavioral state, movement signals from the fCO obviously trigger or release centrally generated transitions in motoneuron activity, e.g., by affecting central rhythm generating networks driving trochanteral motoneuron pools. This conclusion was tested by stimulating the fCO in “fictive rhythmic” preparations, activated by the muscarinic agonist pilocarpine in the otherwise isolated and deafferented mesothoracic ganglion. In this situation, sensory signals from the fCO did in fact reset and entrain rhythmic activity in trochanteral motoneurons. The results indicate for the first time that when the stick insect locomotor system is active, sensory signals from the proprioceptor of one leg joint, i.e., the fCO, pattern motor activity in an adjacent leg joint, i.e., the CT joint, by affecting the central rhythm generating network driving the motoneurons of the adjacent joint.


1986 ◽  
Vol 120 (1) ◽  
pp. 59-77
Author(s):  
ANN CLOAREC

The role of vision in distance, position and size discrimination in prey capture has been investigated in normal adult water stick insects (Ranatra linearis L.: Heteroptera) and in ones with one eye covered. Both monocular and intact Ranatra were able to discriminate between two targets subtending the same angle but presented at different distances. They usually chose the target nearer to their foreleg claws. Although monocular subjects undershot more often than controls, they could still estimate distance correctly. When presented with two different-sized targets at the same distance, both monocular and intact subjects usually preferred the larger target within a 1°-10° range, even though monocular animals chose the larger object less consistently. They were able to distinguish between two targets differing in size by only 1°. Asymmetrical presentations of two identical targets stressed the importance of the central position. Intact animals always preferred the target nearer their midline. These data also revealed the unexpected ability of Ranatra to strike accurately at two targets or prey items simultaneously. When two identical targets were presented simultaneously and symmetrically, aims were directed at both targets, and one was grasped by each raptorial foreleg, thus indicating an absence of confusion.


1985 ◽  
Vol 116 (1) ◽  
pp. 301-311 ◽  
Author(s):  
ULRICH BÄSSLER ◽  
EVA FOTH ◽  
GERHARD BREUTEL

On a slippery surface the forelegs of a decapitated stick insect walk forwards and the hindlegs, backwards. Animals with only forelegs but that are otherwise intact walk forwards, whereas animals with only hindlegs walk mostly backwards. Usually when intact animals start to walk, their hindlegs exert a rearwards thrust on the substrate, but occasionally the starting forces are directed forwards. A rampwise extension of the femoral chordotonal organ in the fixed foreleg of a walking animal first excites the flexor tibiae muscle (positive feedback). Towards the end of the ramp stimulus the activity of the flexor decreases, and the extensor tibiae motor neurones become strongly active. All experiments indicated that the inherent direction of movement of the metathorax is rearwards. In intact animals there must be a coordinating pathway from the prothorax to the metathorax that, together with the suboesophageal ganglion, induces the hindlegs to walk forwards.


1992 ◽  
Vol 173 (1) ◽  
pp. 91-108 ◽  
Author(s):  
R. Kittmann ◽  
J. Schmitz

The femoral chordotonal organ (fCO), one of the largest proprioceptive sense organs in the leg of the stick insect, is important for the control of the femur-tibia joint during standing and walking. It consists of a ventral scoloparium with about 80 sensory cells and a dorsal scoloparium with about 420 sensory cells. The present study examines the function of these scoloparia in the femur-tibia control loop. Both scoloparia were stimulated independently and the responses in the extensor tibiae motoneurones were recorded extra- and intracellularly. The ventral scoloparium, which is the smaller of the two, functions as the transducer of the femur-tibia control loop. Its sensory cells can generate the known resistance reflexes. The dorsal scoloparium serves no function in the femur-tibia control loop and its stimulation elicited no or only minor reactions in the extensor motoneurones. A comparison with other insect leg proprioceptors shows that a morphological subdivision of these organs often indicates a functional specialization.


1989 ◽  
Vol 144 (1) ◽  
pp. 81-111 ◽  
Author(s):  
ANSGAR BÜSCHGES

The femoral chordotonal organ (ChO) of the right middle leg of the inactive stick insect Carausius morosus was stimulated by applying movements having a ramp-like time course, while recordings were made from local and interganglionic interneurones in the anterior ventral median part of the ganglion. Position, velocity and acceleration of the movements were varied independently and the interneurones were categorized on the basis of their responses to the changes in these parameters. Position-sensitivity was always accompanied by responses to velocity and/or acceleration. Velocity-sensitive responses were excitatory or inhibitory and were produced by elongation or relaxation, or by both. In some cases, velocity-sensitive neurones were also affected by position and acceleration. Acceleration responses were always excitatory and were often found in neurones that showed no effects of velocity or position. It is inferred that sensory input from different receptors in the ChO is processed by single interneurones. No interneurone in the recording region was found to be directly involved in the resistance reflex of the extensor tibiae motoneurones, elicited by stimulation of the ChO.


Insects ◽  
2020 ◽  
Vol 11 (1) ◽  
pp. 40 ◽  
Author(s):  
Johannes Strauß

Mechanosensory organs in legs play are crucial receptors in the feedback control of walking and in the detection of substrate-borne vibrations. Stick insects serve as a model for the physiological role of chordotonal organs and campaniform sensilla. This study documents, by axonal tracing, the neural innervation of the complex chordotonal organs and groups of campaniform sensilla in the proximal tibia of the midleg in Sipyloidea sipylus. In total, 6 nerve branches innervate the different sensory structures, and the innervation pattern associates different sensilla types by their position. Sensilla on the anterior and posterior tibia are innervated from distinct nerve branches. In addition, the variation in innervation is studied for five anatomical branching points. The most common variation is the innervation of the subgenual organ sensilla by two nerve branches rather than a single one. The fusion of commonly separated nerve branches also occurred. However, a common innervation pattern can be demonstrated, which is found in >75% of preparations. The variation did not include crossings of nerves between the anterior and posterior side of the leg. The study corrects the innervation of the posterior subgenual organ reported previously. The sensory neuroanatomy and innervation pattern can guide further physiological studies of mechanoreceptor organs and allow evolutionary comparisons to related insect groups.


1995 ◽  
Vol 73 (5) ◽  
pp. 1843-1860 ◽  
Author(s):  
A. Buschges ◽  
H. Wolf

1. Locusts (Locusta migratoria) and stick insects (Carausius morosus) exhibit different strategies for predator avoidance. Locusts rely primarily on walking and jumping to evade predators, whereas stick insects become cataleptic, catalepsy forming a major component of the twig mimesis exhibited by this species. The neuronal networks that control postural leg movements in locusts and stick insects are tuned differently to their specific behavioral tasks. An important prerequisite for the production of catalepsy in the stick insect is the marked velocity dependency of the control network, which appears to be generated at the level of nonspiking local interneurons. We examined interneuronal pathways in the network controlling the femur-tibia joint of the locust middle leg and compared its properties with those described for the stick insect middle leg. It was our aim to identify possible neural correlates of the species-specific behavior with regard to postural leg motor control. 2. We obtained evidence that the neuronal networks that control the femur-tibia joints in the two species consist of morphologically and physiologically similar--and thus probably homologous--interneurons. Qualitatively, these interneurons receive the same input from the femoral chordotonal organ receptors and they drive the same pools of leg motoneurons in both species. 3. Pathways that contribute to the control of the femur-tibia joint include interneurons that support both "resisting" and "assisting" responses with respect to the motoneuron activity that is actually elicited during reflex movements. Signal processing via parallel, antagonistic pathways therefore appears to be a common principle in insect leg motor control. 4. Differences between the two insect species were found with regard to the processing of velocity information provided by the femoral chordotonal organ. Interneuronal pathways are sensitive to stimulus velocity in both species. However, in the locust there is no marked velocity dependency of the interneuronal responses, whereas in the same interneurons of the stick insect it is pronounced. This characteristic was maintained at the level of the motoneurons controlling the femur-tibia joint. Pathways for postural leg motor control in the locust thus lack an important prerequisite for the generation of catalepsy, that is, a marked velocity dependency.


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