The neural basis of catalepsy in the stick insect cuniculina impigra

The femur-tibia control system which is responsible for catalepsy is studied in the open-loop configuration (input: stimulation of the femoral chordotonal organ; output: spike-frequency of FETi and SETi as well as the force produced by the extensor tibiae muscle). Comparison of motor neuron activities and muscle force reveals the input-output relationships of the extensor tibiae muscle. This muscle behaves like a low-pass filter with a small time constant for rising inputs and a large time constant for falling inputs. It forms the decisive low-pass filter for force production of the complete system. For freely moving tibia, the elastic properties of the muscles combined with the inert mass of the tibia contribute to the low-pass filter properties. The muscle does not contribute to the high-pass filter properties of the complete system. During repetitive stimulation FETi habituates quickly.Supported by DFG Ba 578     

The neural basis of catalepsy in the stick insect cuniculina impigra

Intracellular records from somata of FETi and SETi were performed during sinusoidal and rampwise stimulation of the femoral chordotonal organ. The responses were larger on average in SETi but this could be random. All other parameters (form of amplitude-frequency plot, phase-frequency plot, half-lives of rise and fall during ramp-wise stimulations) were not significantly different for both neurons. The responses of both neurons are symmetrical for stretch and release stimuli. Thus, these motor neurons are the decisive rectifiers for the extensor part of the femurtibia control loop. The habituation of FETi during repetitive stimulation is produced by a decrease in response amplitude together with a hyperpolarizing dc-shift. The imput of both neurons behaves like the output of a lead-lag-system with a predominant phasic component, whose high-pass filter has a time constant which depends on input slope. The nonlinear high-pass filter properties which are one important cause of catalepsy can thus be attributed to properties of interneurons and/or chordotonal organ. For most parts of the simulation of the femur-tibia control system (Cruse and Storrer, 1977) the anatomical and physiological correlates could be shown (Fig. 13).     

Systemanalytische Untersuchung eines aufgeschnittenen Regelkreises,der die Beinstellung der Stabheuschrecke Carausius morosus kontrolliert: Kraftmessungen an den Antagonisten Flexor und Extensor tibiae

In the leg of the stick insect Carausius morosus there exists a feedback loop which controls the position of the femur-tibia joint (Bässler, 1965). This feedback mechanism is broken to investigate the open loop system. As output the forces of the two antagonistic muscles flexor tibiae and extensor tibiae are measured separately. As input the feedback transducer of the control mechanism, a chordotonal organ, is stimulated by different kinds of input functions: sine-, step-, delta-, rectangular-and ramp functions. As a qualitative result one can say, that both the flexor-system and the extensor-system have rectifying and high-pass filter properties. However, the comparison between responses to different input functions show that the quantitative properties of this high-pass filter change very strongly with the shape of the input function. Therefore the existence of different nonlinearities has to be assumed.

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Mit Unterstützung der Deutschen Forschungsgemeinschaft (Nr. Ba 578/1)

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Die Arbeit enthält einen Teil der Dissertation von J. Storrer     

Distributed processing on the basis of parallel and antagonistic pathways simulation of the femur-tibia control system in the stick insect

In inactive stick insects, sensory information from the femoral chordotonal organ (fCO) about position and movement of the femur-tibia joint is transferred via local nonspiking interneurons onto extensor and flexor tibiae motoneurons. Information is processed by the interaction of antagonistic parallel pathways at two levels: (1) at the input side of the nonspiking interneurons and (2) at the input side of the motoneurons. We tested by a combination of physiological experiments and computer simulation whether the known network topology and the properties of its elements are sufficient to explain the generation of the motor output in response to passive joint movements, that is resistance reflexes. In reinvestigating the quantitative characteristics of interneuronal pathways we identified 10 distinct types of nonspiking interneurons. Synaptic inputs from fCO afferents onto these interneurons are direct excitatory and indirect inhibitory. These connections were investigated with respect to position and velocity signals from the fCO. The results were introduced in the network simulation. The motor output of the simulation has the same characteristics as the real system, even when particular types of interneurons were removed in the simulation and the real system.     

Physiology of vibration-sensitive afferents in the femoral chordotonal organ of the stick insect

The femoral chordotonal organ in orthopterans signals proprioceptive sensory information concerning the femur-tibia joint to the central nervous system. In the stick insect, 80 out of 500 afferents sense tibial position, velocity, or acceleration. It has been assumed that the other sensory cells in the chordotonal organ would serve as vibration detectors. Extracellular recordings from the femoral chordotonal organ nerve in fact revealed a sensitivity of the sense organ for vibrations with frequencies ranging from 10 Hz to 4 kHz, with a maximum sensitivity between 200 and 800 Hz. Single vibration-sensitive afferents responded to the same range of frequencies. Their spike activity depended on acceleration amplitude and displacement amplitude of the vibration stimulus. Additionally, 80% of the vibration-sensitive afferents received indirect presynaptic inputs from themselves or from other afferents of the femoral chordotonal organ, the amplitude of which depended on stimulus frequency and displacement amplitude. They were associated with a decrease of input resistance in the afferent terminal. From the present investigation we conclude that the femoral chordotonal organ of the stick insect is a bifunctional sensory organ that, on the one hand, measures position and movement of the tibia and, on the other hand, detects vibration of the tibia. Accepted: 6 November 1998     

The network producing the “active reaction” of stick insects is a functional element of different pattern generating systems

Elongation of the femoral chordotonal organ (signalling a flexion movement of the femur-tibia joint) in stick insects being active releases the active reaction (AR) in the extensor and flexor motor neurones. The AR was released in hindlegs in a situation where free animals would preferentially walk backwards. In most cases the coordination between extensor-flexor and the retractor unguis muscle was like in a stance phase of backward walking. In a situation where free animals would preferentially walk forwards, the percentage of ARs was smaller, and resistance reflexes became more frequent. When campaniform sensilla of the hind leg were destroyed coordinations like in a swing phase of forward walking became more frequent. — Additional stimuli during searching movements in an artificially closed femur-tibia feedback system (Weiland et al. 1986) showed that the AR is expressed also under these conditions and controls velocity and endpoint of a flexion movement. All results support the idea that the neural system producing the AR is a functional element of the pattern generator for forward walking, of the one for backward walking and of the one for searching movements. As far as this system is concerned the three pattern generators only differ in the kind of coordinating pathways between constant functional elements.     

Time series modeling of neuromuscular system

The dynamic response of the human ankle joint to a bandlimited random torque perturbation superimposed on a constant bias torque is observed in normal human subjects. The applied torque input, the joint angular rotation output, and the electromyographic activity using surface electrodes from the extensor and the flexor muscles of the ankle joint were recorded. Transfer function models using time series techniques were developed for the torque — angular rotation input-output pair and for the angular rotation — electromyographic activity input-output pair. A parameter constraining technique was applied to develop more reliable models. It is shown that the asymptotic behavior of the system must be taken into account during parameter optimization to develop better predictive models.This work was supported in part by National Science Foundation grant ENG-7608754 and grants from the National Institutes of Health NS-12877 and NS-00196     

Figure-ground discrimination by relative movement in the visual system of the fly

The visual system of the fly performs various computations on photoreceptor outputs. The detection and measurement of movement is based on simple nonlinear multiplication-like interactions between adjacent pairs and groups of photoreceptors. The position of a small contrasted object against a uniform background is measured, at least in part, by (formally) 1-input nonlinear flicker detectors. A fly can also detect and discriminate a figure that moves relative to a ground texture. This computation of relative movement relies on a more complex algorithm, one which detects discontinuities in the movement field. The experiments described in this paper indicate that the outputs of neighbouring movement detectors interact in a multiplication-like fashion and then in turn inhibit locally the flicker detectors. The following main characteristic properties (partly a direct consequence of the algorithm's structure) have been established experimentally: a) Coherent motion of figure and ground inhibit the position detectors whereas incoherent motion fails to produce inhibition near the edges of the moving figure (provided the textures of figure and ground are similar). b) The movement detectors underlying this particular computation are direction-insensitive at input frequencies (at the photoreceptor level) above 2.3 Hz. They become increasingly direction-sensitive for lower input frequencies. c) At higher input frequencies the fly cannot discriminate an object against a texture oscillating at the same frequency and amplitude at 0° and 180° phase, whereas 90° or 270° phase shift between figure and ground oscillations yields maximum discrimination. d) Under conditions of coherent movement, strong spatial incoherence is detected by the same mechanism. The algorithm underlying the relative movement computation is further discussed as an example of a coherence measuring process, operating on the outputs of an array of movement detectors. Possible neural correlates are also mentioned.     

Adaptive control for insect leg position: controller properties depend on substrate compliance

This paper concentrates on the system that controls the femur-tibia joint in the legs of the stick insect, Carausius morosus. Earlier investigations have shown that this joint is subject to a mixture of proportional and differential control whereby the differential part plays a prominent role. Experiments presented here suggest another interpretation: single legs of a stick insect were systematically perturbed using devices of different compliance and compensatory forces and movements monitored. When the compliance is high (soft spring), forces are generated that return the leg close to its original position. When the compliance is low (stiff spring), larger forces are generated but sustained changes in position occur that are proportional to the force that is applied. Selective ablation of leg sense organs showed that the leg did not maintain its position after elimination of afferents of the femoral chordotonal organ. Ablation of leg campaniform sensilla had no effect. These data support the idea that different control strategies are used, depending upon substrate compliance. In particular, what we and other authors have called a differential controller, is now considered as an integral controller that intelligently gives up when the correlation between motor output and movement of the leg is low.We would like to dedicate this article to Prof. Dr. Ulrich Bässler. Starting in the 1960s, his seminal work stimulated a long series of fruitful studies that, even today, reveal exciting insights into motor control.     

Sensory influences on the coordination of two leg joints during searching movements of stick insects

Animals (Cuniculina impigra) possessing only one foreleg with restrained coxa perform very stereotyped searching movements during which the movements of the femur-tibia and coxa-trochanter joints are well coordinated. After ablation of either hairfield BF1 (measuring the position of the coxa-trochanter joint) or the apodeme of the femoral chordotonal organ (measuring the position of the femur-tibia joint) each joint can still be moved but the coordination changes and becomes very labile. The consequences for the ideas about the construction principles of the pattern generator for searching movements are discussed.     

Properties of the feedback system controlling the coxa-trochanter joint in the stick insect Carausius morosus

The angle of the coxa-trochanter (C-T) joint in the stick insect Carausius morosus is controlled by a negative feedback mechanism. It is shown that the trochanteral hair plate alone functions as the feedback transducer and that the rhomboid hair plate is not involved in the feedback loop.The properties of the C-T control system were investigated by means of force measurements. The results cannot be adequately described in all details by either a fractional differentiator model, a model which fits many sensory systems, or a nonlinear bandpass filter, a model which fits the force response of the femur-tibia feedback loop. The fractional differentiator model adequately describes the frequency response of the open-loop system to sinusoidal stimulation with 34 deg stimulus amplitude. However, the responses to sinusoidal and steplike stimulation with 10 deg stimulus amplitude do not fit this model. They are better described by the model of a nonlinear bandpass filter.The possible contribution of mechanical properties of the musculature and the joint to the total force response is discussed. It is suggested that cocontractions occurring at higher stimulus frequencies alters the muscle properties and enables the animal to respond to stimulus frequencies above the upper corner frequency of the active feedback loop.     

Sensorimotor pathways involved in interjoint reflex action of an insect leg

Coordination of motor output between leg joints is crucial for the generation of posture and active movements in multijointed appendages of legged organisms. We investigated in the stick insect the information flow between the middle leg femoral chordotonal organ (fCO), which measures position and movement in the femur-tibia (FT) joint and the motoneuron pools supplying the next proximal leg joint, the coxa-trochanteral (CT) joint. In the inactive animal, elongation of the fCO (by flexing the FT joint) induced a depolarization in eight of nine levator trochanteris motoneurons, with a suprathreshold activation of one to three motoneurons. Motoneurons of the depressor trochanteris muscle were inhibited by fCO elongation. Relaxation signals, i.e., extension of the FT joint, activated both levator and depressor motoneurons; i.e., both antagonistic muscles were coactivated. Monosynaptic as well as polysynaptic pathways contribute to interjoint reflex actions in the stick insect leg. fCO afferents were found to induce short latency EPSPs in levator motoneurons, providing evidence for direct connections between fCO afferents and levator motoneurons. In addition, neuronal pathways via intercalated interneurons were identified that transmit sensory information from the fCO onto levator and/or depressor motoneurons. Finally, we describe two kinds of alterations in interjoint reflex action: (a) With repetitive sensory stimulation, this interjoint reflex action shows a habituation-like decrease in strength. (b) In the actively moving animal, interjoint reflex action in response to fCO elongation, mimicking joint flexion, qualitatively remained the same sign, but with a marked increase in strength, indicating an increased influence of sensory signals from the FT joint onto the adjacent CT joint in the active animal. © 1997 John Wiley & Sons, Inc. J Neurobiol 33: 891–913, 1997     

A model of anuran retina relating interneurons to ganglion cell responses

A model is presented which accounts for many characteristic response properties used to classify anuran ganglion cell types while being consistent with data concerning interneurons. In the model color is ignored and input stimuli are assumed to be only black and white at high contrast. We show that accurate ganglion cell responses are obtained even with simplified receptors and horizontal cells: Receptors are modeled as responding with a step change, while horizontal cells respond only to global changes in intensity brought about by full field illumination changes. A hyperpolarizing and depolarizing bipolar cell are generated y subtracting local receptor and horizontal potentials. Two transient amacrine cells (On and Off) are generated using a high-pass filter like mechanism with a thresholded output which responds to positive going changes in the corresponding bipolar cell potentials. The model shows how a selective combination of bipolar and amacrine channels can account for many of the response properties used to classify the anuran ganglion cell types (class-0 through 4) and makes several experimental predictions.     

Dynamic response properties of movement detectors: Theoretical analysis and electrophysiological investigation in the visual system of the fly

Dynamic aspects of the computation of visual motion information are analysed both theoretically and experimentally. The theoretical analysis is based on the type of movement detector which has been proposed to be realized in the visual system of insects (e.g. Hassenstein and Reichardt 1956; Reichardt 1957, 1961; Buchner 1984), but also of man (e.g. van Doorn and Koenderink 1982a, b; van Santen and Sperling 1984; Wilson 1985). The output of both a single movement detector and a one-dimensional array of detectors is formulated mathematically as a function of time. The resulting movement detector theory can be applied to a much wider range of moving stimuli than has been possible on the basis of previous formulations of the detector output. These stimuli comprise one-dimensional smooth detector input functions, i.e. functions which can be expanded into a time-dependent convergent Taylor series for any value of the spatial coordinate.The movement detector response can be represented by a power series. Each term of this series consists of one exclusively time-dependent component and of another component that depends, in addition, on the properties of the pattern. Even the exclusively time-dependent components of the movement detector output are not solely determined by the stimulus velocity. They rather depend in a non-linear way on the weighted sum of the instantaneous velocity and all its higher order time derivatives. The latter point represents another reason — not discussed so far in the literature — that movement detectors of the type analysed here do not represent pure velocity sensors.The significance of this movement detector theory is established for the visual system of the fly. This is done by comparing the spatially integrated movement detector response with the functional properties of the directionally-selective motion-sensitive. Horizontal Cells of the third visual ganglion of the fly's brain.These integrate local motion information over large parts of the visual field. The time course of the spatially integrated movement detector response is about proportional to the velocity of the stimulus pattern only as long as the pattern velocity and its time derivatives are sufficiently small. For large velocities and velocity changes of the stimulus pattern characteristic deviations of the response profiles from being proportional to pattern velocity are predicted on the basis of the detector theory developed here. These deviations are clearly reflected in the response of the wide-field Horizontal Cells, thus, providing very specific evidence that the movement detector theory developed here can be applied to motion detection in the fly. The characteristic dynamic features of the theoretically predicted and the experimentally determined cellular responses are exploited to estimate the time constant of the movement detector filter.     

Gain control in a proprioceptive feedback loop as a prerequisite for working close to instability

In the artificially closed femur-tibia control system of stick insects oscillations were induced in 3 different ways: Increasing the phase-shift by introducing an electronic delay, afference sign reversal and coupling the tibia to an inert mass. In all 3 cases the oscillations stopped after some time. The gain of the open-loop system was significantly smaller after the oscillations. Afference sign reversal by surgically crossing of the receptor apodeme of the femoral chordotonal organ for 25–85 days does not lead to altered characteristics of the control loop. When sinusoidal passive movements are forced upon the intact femur-tibia joint the forces resisting these movements do not decrease with time. In contrast to direct stimulation of the femoral chordotonal organ, these passive movements also influence the contralateral leg. The experiments show that the gain-control system of the femur-tibia control loop of stick insects consists of at least two components: A sensitization system (with inputs from many kinds of stimuli indicating some kind of disturbance) increases the gain of all reflex loops. A specific habituation-like system decreases the gain with repetitive stimulation only of one control system.Abbreviations fCO femoral chordotonal organ - SETi slow extensor tibiae motor neuron     

Open loop analysis of a feedback mechanism controlling the leg position in the stick insect Carausius morosus: Comparison between experiment and simulation

In the legs of the stick insect Carausius morosus a feedback mechanism exists to control the value of the angle between femur and tibia. It is possible to investigate the open loop system by moving as input experimentally the receptor apodeme of the femoral chordotonal organ, which acts as feedback transducer measuring the angle between femur and tibia (Bässler, 1965). As output the forces are measured separately which are developed by the two antagonistic muscles moving the femur-tibia joint. The response of this system to different step-, sine-, ramp-and -functions are measured. An electronic analog model is constructed to simulate the biological system (Fig. 1). Although a number of different nonlinearities arise in the biological system, as a first-order approximation the model shows a sufficient fit to the experimental results (Figs. 2–9). The main characteristics of the model are as follows. It consists of two independent subsystems, the flexor system and the extensor system. Each subsystem again consists of two parallel branches with high-pass properties of different time constants. In each subsystem one branch is only excitable by input functions of a slope smaller than a certain degree. It is remarkable, that no mutual inhibitory influence between the sub-systems controlling the antagonistic muscles is necessary in the model.Supported by the Deutsche Forschungsgemeinschaft (Nr. Ba 578/1)     

A computational simulated control system for a high-force pneumatic muscle actuator: system definition and application as an augmented orthosis

High-force pneumatic muscle actuators (PMAs) are used for force assistance with minimal displacement applications. However, poor control due to dynamic nonlinearities has limited PMA applications. A simulated control system is developed consisting of: (1) a controller relating an input position angle to an output proportional pressure regulator voltage, (2) a phenomenological model of the PMA with an internal dynamic force loop (system time constant information), (3) a physical model of a human sit-to-stand task and (4) an external position angle feed-back loop. The results indicate that PMA assistance regarding the human sit-to-stand task is feasible within a specified PMA operational pressure range.     

Coding proprioceptive information to control movement to a target: Simulation with a simple neural network

When the stick insect walks, the middle and rear legs step to positions immediately behind the tarsus of the adjacent rostral leg. Previous reports have described this movement to a target as a relationship between the tarsus positions of the two legs in a Cartesian coordinate system. However, leg proprioceptors measure the position of the target leg in terms of joint angles and leg muscles bring the tarsus of the moving leg to the proper end-point by establishing appropriate angles at the joints. Representation of this task in Cartesian coordinates requires non-linear coordinate transformations; realizing such a transformation in the nervous system appears to require many neurons. The present simulation using the back-propagation algorithm shows that a simple network of only nine units — 3 sensory input units, 3 motor output units, and 3 hidden units — suffices. The simulation also shows that an analytic coordinate transformation can be replaced by a direct association of joint configurations in the moving leg with those in the target leg.     

A computational simulated control system for a high-force pneumatic muscle actuator: system definition and application as an augmented orthosis

High-force pneumatic muscle actuators (PMAs) are used for force assistance with minimal displacement applications. However, poor control due to dynamic nonlinearities has limited PMA applications. A simulated control system is developed consisting of: (1) a controller relating an input position angle to an output proportional pressure regulator voltage, (2) a phenomenological model of the PMA with an internal dynamic force loop (system time constant information), (3) a physical model of a human sit-to-stand task and (4) an external position angle feed-back loop. The results indicate that PMA assistance regarding the human sit-to-stand task is feasible within a specified PMA operational pressure range.     

Static and dynamic human flexor tendon–pulley interaction

The aim of the study was to investigate the influence of a preceding flexion or extension movement on the static interaction of human finger flexor tendons and pulleys concerning flexion torque being generated. Six human fresh frozen cadaver long fingers were mounted in an isokinetic movement device for the proximal interphalangeal (PIP) joint. During flexion and extension movement both flexor tendons were equally loaded with 40 N while the generated moment was depicted simultaneously at the fingertip. The movement was stopped at various positions of the proximal interphalangeal joint to record dynamic and static torque. The static torque was always greater after a preceding extension movement compared to a preceding flexion movement in the corresponding same position of the joint. This applied for the whole arc of movement of 0–105°. The difference between static extension and flexion torque was maximal 11% in average at about 83° of flexion. Static torque was always smaller than dynamic torque during extension movement and always greater than dynamic torque during flexion movement. The kind of preceding movement therefore showed an influence to the torque being generated in the proximal interphalangeal joint. The effect could be simulated on a mechanical finger device.