Biomedical subjects

Volker Dürr

Publications and source records attributed to Volker Dürr.

9 recordsLinked to original sources

PubMed2007

Insect walking is based on a decentralized architecture revealing a simple and robust controller.

Control of walking in rugged terrain requires one to incorporate different issues, such as the mechanical properties of legs and muscles, the neuronal control structures for the single leg, the mechanics and neuronal control structures for the coordination between legs, as well as central decisions that are based on external information and on internal states. Walking in predictable environments and fast running, to a large degree, rely on muscle mechanics. Conversely, slow walking in unpredictable terrain, e.g. climbing in rugged structures, has to rely on neuronal systems that monitor and intelligently react to specific properties of the environment. An arthropod model system that shows the latter abilities is the stick insect, based on which this review will be focused. An insect, when moving its six legs, has to control 18 joints, three per leg, and therefore has to control 18 degrees of freedom (d.f.). As the body position in space is determined by 6 d.f. only, there are 12 d.f. open to be selected. Therefore, a fundamental problem is as to how these extra d.f. are controlled. Based mainly on behavioural experiments and simulation studies, but also including neurophysiological results, the following control structures have been revealed. Legs act as basically independent systems. The quasi-rhythmic movement of the individual leg can be described to result from a structure that exploits mechanical coupling of the legs via the ground and the body. Furthermore, neuronally mediated influences act locally between neighbouring legs, leading to the emergence of insect-type gaits. The underlying controller can be described as a free gait controller. Cooperation of the legs being in stance mode is assumed to be based on mechanical coupling plus local positive feedback controllers. These controllers, acting on individual leg joints, transform a passive displacement of a joint into an active movement, generating synergistic assistance reflexes in all mechanically coupled joints. This architecture is summarized in the form of the artificial neural network, Walknet, that is heavily dependent on sensory feedback at the proprioceptive level. Exteroceptive feedback is exploited for global decisions, such as the walking direction and velocity.

Animals↗

PubMed2006

Slanted joint axes of the stick insect antenna: an adaptation to tactile acuity.

Like many flightless, obligatory walking insects, the stick insect Carausius morosus makes intensive use of active antennal movements for tactile near range exploration and orientation. The antennal joints of C. morosus have a peculiar oblique and non-orthogonal joint axis arrangement. Moreover, this arrangement is known to differ from that in crickets (Ensifera), locusts (Caelifera) and cockroaches (Blattodea), all of which have an orthogonal joint axis arrangement. Our hypothesis was that the situation found in C. morosus represents an important evolutionary trait of the order of stick and leaf insects (Phasmatodea). If this was true, it should be common to other species of the Phasmatodea. The objective of this comparative study was to resolve this question. We have measured the joint axis orientation of the head-scape and scape-pedicel joints along with other parameters that affect the tactile efficiency of the antenna. The obtained result was a complete kinematic description of the antenna. This was used to determine the size and location of kinematic out-of-reach zones, which are indicators of tactile acuity. We show that the oblique and non-orthogonal arrangement is common to eight species from six sub-families indicating that it is a synapomorphic character of the Euphasmatodea. This character can improve tactile acuity compared to the situation in crickets, locusts and cockroaches. Finally, because molecular data of a recent study indicate that the Phasmatodea may have evolved as flightless, obligatory walkers, we argue that the antennal joint axis arrangement of the Euphasmatodea reflects an evolutionary adaptation to tactile near range exploration during terrestrial locomotion.

Animals↗

PubMed2006

Co-contraction and passive forces facilitate load compensation of aimed limb movements.

Vertebrates and arthropods are both capable of load compensation during aimed limb movements, such as reaching and grooming. We measured the kinematics and activity of individual motoneurons in loaded and unloaded leg movements in an insect. To evaluate the role of active and passive musculoskeletal properties in aiming and load compensation, we used a neuromechanical model of the femur-tibia joint that transformed measured extensor and flexor motoneuron spikes into joint kinematics. The model comprises three steps: first, an activation dynamics module that determines the time course of isometric force; second, a pair of antagonistic muscle models that determine the joint torque; and third, a forward dynamics simulation that calculates the movement of the limb. The muscles were modeled in five variants, differing in the presence or absence of force-length-velocity characteristics of the contractile element, a parallel passive elastic element, and passive joint damping. Each variant was optimized to yield the best simulation of measured behavior. Passive muscle force and viscous joint damping were sufficient and necessary to simulate the observed movements. Elastic or damping properties of the active contractile element could not replace passive elements. Passive elastic forces were similar in magnitude to active forces caused by muscle contraction, generating substantial joint stiffness. Antagonistic muscles co-contract, although there was no motoneuronal coactivation, because of slow dynamics of muscle activation. We quantified how co-contraction simplified load compensation by demonstrating that a small variation of the motoneuronal input caused a large change in joint torque.

Animals↗

PubMed2006

Perturbation of leg protraction causes context-dependent modulation of inter-leg coordination, but not of avoidance reflexes.

All animals capable of legged locomotion execute fast, adaptive compensatory movements in response to perturbation of a step cycle. In terms of motor control, such adaptive behaviour typically involves changes in the kinematics of the perturbed limb as well as changes in coordination between legs. Moreover, the unpredictable variety of real life situations implies that compensatory responses should be sensitive to the behavioural context of the animal. We have investigated the extent to which the compensatory response of a walking stick insect (Carausius morosus) adapts in parallel to strong context-dependent adaptation of step kinematics and inter-leg coordination. The behavioural contexts we chose were straight walking and visually induced curve walking, for both of which the steady state limb kinematics and inter-leg coupling strengths were known. In case of curve walking, we further distinguished contexts according to whether the inner or the outer leg was perturbed. The three contexts differed strongly with respect to the set of joint actions before perturbation. Upon mechanical perturbation of front leg protraction, we studied context-dependent differences in a local avoidance reflex of the perturbed leg, as well as in coordination mechanisms that couple the step cycles of the perturbed leg to its unperturbed neighbours. In all three walking contexts, obstacle contact caused an avoidance movement of the front leg that deviated from the unperturbed swing trajectory. Swing duration was increased while step distance was decreased; however, both effects vanished in the subsequent unperturbed step. The prevailing immediate reaction of the three leg joints were retraction of the coxa (>76%), levation of the femur (>80%), and flexion of the tibia (>80%), regardless of the behavioural context and, therefore, joint action prior to perturbation. Moreover, activation of each one of these joint actions was shown to be independent of the other two. Thus, local avoidance reflexes are not modulated by the descending visual information that causes transition from straight to curve walking, but are composed of context-independent joint actions. Perturbation of the front leg also caused significant shifts of the touch-down position of the perturbed leg and of its unperturbed neighbours. If the inner front leg was perturbed, this shift could persist until the subsequent step. Perturbation affected both the spatial location and the timing of touch-down and lift-off transitions in unperturbed neighbouring legs. These effects on inter-leg coordination were context-dependent. For example, time delay to lift-off of the contralateral neighbour was shortened in inner and straight walking legs, but not in outer legs. Finally, a targeting mechanism that determines foot placement in stick insects was shown to be affected by perturbation in a context-dependent manner. We conclude that the immediate compensatory response of the perturbed leg is not adapted to the behavioural context in spite of strongly differing step kinematics, whereas the compensatory effect on inter-limb coupling is context-dependent.

Animals↗

PubMed2005

The behavioural transition from straight to curve walking: kinetics of leg movement parameters and the initiation of turning.

The control of locomotion requires the ability to adapt movement sequences to the behavioural context of the animal. In hexapod walking, adaptive behavioural transitions require orchestration of at least 18 leg joints and twice as many muscle groups. Although kinematics of locomotion has been studied in several arthropod species and in a range of different behaviours, almost nothing is known about the transition from one behavioural state to another. Implicitly, most studies on context-dependency assume that all parameters that undergo a change during a behavioural transition do so at the same rate. The present study tests this assumption by analysing the sequence of kinematic events during turning of the stick insect Carausius morosus, and by measuring how the time courses of the changing parameters differ between legs. Turning was triggered reliably at a known instant in time by means of the optomotor response to large-field visual motion. Thus, knowing the start point of the transition, the kinematic parameters that initiate turning could be ranked according to their time constants. Kinematics of stick insect walking vary considerably among trials and within trials. As a consequence, the behavioural states of straight walking and curve walking are described by the distributions of 13 kinematic parameters per leg and of orientation angles of head and antennae. The transitions between the behavioural states are then characterised by the fraction of the variance within states by which these distributions differ, and by the rate of change of the corresponding time courses. The antennal optomotor response leads that of the locomotor system. Visually elicited turning is shown to be initiated by stance direction changes of both front legs. The transition from straight to curve walking in stick insects follows different time courses for different legs, with time constants of kinematic parameters ranging from 1.7 s to more than 3 s. Therefore, turning is a behavioural transition that involves a characteristic orchestration of events rather than synchronous parallel actions with a single time constant.

Animals↗

PubMed2005

Context-dependent changes in strength and efficacy of leg coordination mechanisms.

Appropriate coordination of stepping in adjacent legs is crucial for stable walking. Several leg coordination rules have been derived from behavioural experiments on walking insects, some of which also apply to arthropods with more than six legs and to four-legged walking vertebrates. Three of these rules affect the timing of stance-swing transition [rules 1 to 3 (sensu Cruse)]. They can give rise to normal leg coordination and adaptive responses to disturbances, as shown by kinematic simulations and dynamic hardware tests. In spite of their importance to the study of animal walking, the coupling strength associated with these rules has never been measured experimentally. Generally coupling strength of the underlying mechanisms has been considered constant rather than context-dependent. The present study analyses stepping patterns of the stick insect Carausius morosus during straight and curve walking sequences. To infer strength and efficacy of coupling between pairs of sender and receiver legs, the likelihood of the receiver leg being in swing is determined, given a certain delay relative to the time of a swing-stance (or stance-swing) transition in the sender leg. This is compared to a corresponding measure for independent, hence uncoupled, step sequences. The difference is defined as coupling strength. The ratio of coupling strength and its theoretical maximum is defined as efficacy. Irrespective of the coordination rule, coupling strength between ipsilateral leg pairs is at least twice that of contralateral leg pairs, being strongest between ipsilateral hind and middle legs and weakest between contralateral middle legs. Efficacy is highest for inhibitory rule 1, reaching 84-95% for ipsilateral and 29-65% for contralateral leg pairs. Efficacy of excitatory rules 2 and 3 ranges between 35-56% for ipsilateral and 8-21% for contralateral leg pairs. The behavioural transition from straight to curve walking is associated with context-dependent changes in coupling strength, increasing in both outer leg pairs and decreasing between inner hind and middle leg. Thus, the coordination rules that are thought to underlie many adaptive properties of the walking system, themselves adapt in a context-dependent manner.

Animals↗

PubMed2004

Tactile efficiency of insect antennae with two hinge joints.

Antennae are the main organs of the arthropod tactile sense. In contrast to other senses that are capable of retrieving spatial information, e.g. vision, spatial sampling of tactile information requires active movement of the sense organ. For a quantitative analysis of basic principles of active tactile sensing, we use a generic model of arbitrary antennae with two hinge joints (revolute joints). This kind of antenna is typical for Orthoptera and Phasmatodea, i.e. insect orders that contain model species for the study of antennal movements, including cricket, locust and stick insect. First, we analyse the significance of morphological properties on workspace and sampling acuity. It is shown how joint axis orientation determines areas out of reach while affecting acuity in the areas within reach. Second, we assume a parametric set of movement strategies, based on empirical data on the stick insect Carausius morosus, and investigate the role of each strategy parameter on tactile sampling performance. A stochastic environment is used to measure sampling density, and a viscous friction model is assumed to introduce energy consumption and, thus, a measure of tactile efficiency. Up to a saturation level, sampling density is proportional to the range or frequency of joint angle modulation. The effect of phase shift is strong if joint angle modulation frequencies are equal, but diminishes for other frequency ratios. Speed of forward progression influences the optimal choice of movement strategy. Finally, for an analysis of environmental effects on tactile performance, we show how efficiency depends on predominant edge direction. For example, with slanted and non-orthogonal joint axis orientations, as present in the stick insect, the optimal sampling strategy is less sensitive to a change from horizontal to vertical edge predominance than with orthogonal and non-slanted joint axes, as present in a cricket.

Animals↗

PubMed2004

A posture optimization algorithm for model-based motion capture of movement sequences.

We have developed and evaluated a new optical motion capture approach that is suitable for a wide range of studies in neuroethology and motor control. Based on the stochastic search algorithm of Simulated Annealing (SA), it utilizes a kinematic body model that includes joint angle constraints to reconstruct posture from an arbitrary number of views. Rather than tracking marker trajectories in time, the algorithm minimizes an error function that compares predicted model projections to the recorded views. Thus, each video-frame is analyzed independently from other frames, enabling the system to recover from incorrectly analyzed postures. The system works with standard computer and video equipment. Its accuracy is evaluated using videos of animated locust leg movements, recorded by two orthogonal views. The resulting joint angle RMS errors range between 0.7 degrees and 4.9 degrees, limited by the pixel resolution of the digital video. 3D-movement reconstruction is possible even from a single view. In a real experimental application, stick insect walking sequences are analyzed with leg joint angle deviations between 0.5 degrees and 3.0 degrees. This robust and accurate performance is reached in spite of marker fusions and occlusions, simply by exploiting the natural constraints imposed by a kinematic chain and a known experimental setup.

Algorithms↗

PubMed2003

Load compensation in targeted limb movements of an insect.

The task of a multi-jointed limb making an aimed movement towards a target requires that the movement is regulated against external perturbations such as changing load. In particular, loading one part of a limb leads to altered static forces on all proximal segments, and to additional dynamic joint interaction forces when the limb moves. We have addressed the question of load compensation in an insect preparation in which a locust makes aimed scratching movements with a hind leg in response to tactile stimulation of a wing. We show that loading the femur or tibia with the equivalent of 8.5 times the mass of the tibia (corresponding to an increase of up to 11.6 times the rotational moment of inertia at the femur-tibia joint) does not impair the animal's ability to make well-coordinated, aimed movements of that leg towards different targets. The kinematics of the movements are the same, and animals aim the same part of their distal tibia at the target, regardless of loading. The movements are carried out with equal accuracy and at the same initial velocity under all load conditions. Because loading of the leg does not change the behavioural performance, there is no indication of a change in aiming strategy. This implies high leg joint stiffness and/or the existence of high gain proprioceptive control loops. We have previously shown that in the unloaded condition, movements elicited by stimuli to different places on the wing are driven by a single underlying movement pattern that shifts depending on stimulus location along the wing surface. Our present data show that leg proprioceptive inputs are also integrated into the leg motor networks, rendering hind limb targeting robust against large changes in moment of inertia.

Animals↗