Navigating complex terrains requires dynamic interactions between the substrate, musculoskeletal, and sensorimotor systems. Current perturbation studies have mostly used visible terrain height perturbations, which do not allow us to distinguish among the neuromechanical contributions of feedforward control, feedback-mediated, and mechanical perturbation responses. Here, we use treadmill-belt speed perturbations to induce a targeted perturbation to foot speed only, and without terrain-induced changes in joint posture and leg loading at stance onset. Based on previous studies suggesting a proximo-distal gradient in neuromechanical control, we hypothesized that distal joints would exhibit larger changes in joint kinematics, compared to proximal joints. Additionally, we expected birds to use feedforward strategies to increase the intrinsic stability of gait. To test these hypotheses, seven adult guinea fowl were video recorded while walking on a motorized treadmill, during both steady and perturbed trials. Perturbations consisted of repeated exposures to a deceleration and acceleration of the treadmill-belt speed. Surprisingly, we found that joint angular trajectories and center of mass fluctuations remain very similar, despite substantial perturbation of foot velocity by the treadmill belt. Hip joint angular trajectories exhibit the largest changes, with the birds adopting a slightly more flexed position across all perturbed strides. Additionally, we observed increased stride duration across all strides, consistent with feedforward changes in the control strategy. The speed perturbations mainly influenced the timing of stance and swing, with the largest kinematic changes in the strides directly following a deceleration. Our findings do not support the general hypothesis of a proximo-distal gradient in joint control, as distal joint kinematics remain largely unchanged. Instead, we find that leg angular trajectory and the timing of stance and swing are most sensitive to this specific perturbation, and leg length actuation remains largely unchanged. Our results are consistent with modular task-level control of leg length and leg angle actuation, with different neuromechanical control and perturbation sensitivity in each actuation mode. Distal joints appear to be sensitive to changes in vertical loading but not foot fore-aft velocity. Future directions should include in vivo studies of muscle activation and force–length dynamics to provide more direct evidence of the sensorimotor control strategies for stability in response to belt-speed perturbations.
Limb, joint and pelvic kinematic control in the quail coping with steps upwards and downwards
Small cursorial birds display remarkable walking skills and can negotiate complex and unstructured terrains with ease. The neuromechanical control strategies necessary to adapt to these challenging terrains are still not well understood. Here, we analyzed the 2D- and 3D pelvic and leg kinematic strategies employed by the common quail to negotiate visible steps (upwards and downwards) of about 10%, and 50% of their leg length. We used biplanar fluoroscopy to accurately describe joint positions in three dimensions and performed semi-automatic landmark localization using deep learning. Quails negotiated the vertical obstacles without major problems and rapidly regained steady-state locomotion. When coping with step upwards, the quail mostly adapted the trailing limb to permit the leading leg to step on the elevated substrate similarly as it did during level locomotion. When negotiated steps downwards, both legs showed significant adaptations. For those small and moderate step heights that did not induce aerial running, the quail kept the kinematic pattern of the distal joints largely unchanged during uneven locomotion, and most changes occurred in proximal joints. The hip regulated leg length, while the distal joints maintained the spring-damped limb patterns. However, to negotiate the largest visible steps, more dramatic kinematic alterations were observed. There all joints contributed to leg lengthening/shortening in the trailing leg, and both the trailing and leading legs stepped more vertically and less abducted. In addition, locomotion speed was decreased. We hypothesize a shift from a dynamic walking program to more goal-directed motions that might be focused on maximizing safety.
more » « less- Award ID(s):
- 2015317
- NSF-PAR ID:
- 10371801
- Publisher / Repository:
- Nature Publishing Group
- Date Published:
- Journal Name:
- Scientific Reports
- Volume:
- 12
- Issue:
- 1
- ISSN:
- 2045-2322
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
More Like this
-
more » « less
Abstract -
Intent recognition in lower-limb assistive devices typically relies on neuromechanical sensing of an affected limb acquired through embedded device sensors. It remains unknown whether signals from more widespread sources such as the contralateral leg and torso positively influence intent recognition, and how specific locomotor tasks that place high demands on the neuromuscular system, such as changes of direction, contribute to intent recognition. In this study, we evaluated the performances of signals from varying mechanical modalities (accelerographic, gyroscopic, and joint angles) and locations (the trailing leg, leading leg and torso) during straight walking, changes of direction (cuts), and cuts to stair ascent with varying task anticipation. Biomechanical information from the torso demonstrated poor performance across all conditions. Unilateral (the trailing or leading leg) joint angle data provided the highest accuracy. Surprisingly, neither the fusion of unilateral and torso data nor the combination of multiple signal modalities improved recognition. For these fused modality data, similar trends but with diminished accuracy rates were reported during unanticipated conditions. Finally, for datasets that achieved a relatively accurate (≥90%) recognition of unanticipated tasks, these levels of recognition were achieved after the mid-swing of the trailing/transitioning leg, prior to a subsequent heel strike. These findings suggest that mechanical sensing of the legs and torso for the recognition of straight-line and transient locomotion can be implemented in a relatively flexible manner (i.e., signal modality, and from the leading or trailing legs) and, importantly, suggest that more widespread sensing is not always optimal.more » « less
-
null (Ed.)Abstract Successful motor control requires accurate estimation of our body in space for planning, executing, and evaluating the outcome of our actions. It has been shown that the estimation of limb position is susceptible to motor adaptation. However, a similar effect has not been found in locomotion, possibly due to how it was tested. We hypothesized that split-belt walking with the legs moving at different speeds changes the estimation of the legs’ position when taking a step. Thus, we assessed young subjects’ perception of step length (i.e., inter-feet distance at foot landing) when they moved their legs (active perception) or when the legs were moved by the experimenter (passive perception). We found that the active perception of step length was substantially altered following split-belt walking, whereas passive perception exhibited minor changes. This suggests that split-belt walking induced the adaptation of efferent signals, without altering sensory signals. We also found that active perceptual shifts were sensitive to how they were tested: they were most salient in the trailing leg and at short step lengths. Our results suggest that split-belt walking could modulate the deficient perception of step length post-stroke, which may contribute to gait asymmetries impairing patients’ mobility.more » « less
-
Abstract This paper explores the kinematic synthesis, design, and pilot experimental testing of a six-legged walking robotic platform able to traverse through different terrains. We aim to develop a structured approach to designing the limb morphology using a relaxed kinematic task with incorporated conditions on foot-environments interaction, specifically contact force direction and curvature constraints, related to maintaining contact. The design approach builds up incrementally starting with studying the basic human leg walking trajectory and then defining a “relaxed” kinematic task. The “relaxed” kinematic task consists only of two contact locations (toe-off and heel-strike) with higher-order motion task specifications compatible with foot-terrain(s) contact and curvature constraints in the vicinity of the two contacts. As the next step, an eight-bar leg image is created based on the “relaxed” kinematic task and incorporated within a six-legged walking robot. Pilot experimental tests explore if the proposed approach results in an adaptable behavior which allows the platform to incorporate different walking foot trajectories and gait styles coupled to each environment. The results suggest that the proposed “relaxed” higher-order motion task combined with the leg morphological properties and feet material allowed the platform to walk stably on the different terrains. Here we would like to note that one of the main advantages of the proposed method in comparison with other existing walking platforms is that the proposed robotic platform has carefully designed limb morphology with incorporated conditions on foot-environment interaction. Additionally, while most of the existing multilegged platforms incorporate one actuator per leg, or per joint, our goal is to explore the possibility of using a single actuator to drive all six legs of the platform. This is a critical step which opens the door for the development of future transformative technology that is largely independent of human control and able to learn about the environment through their own sensory systems.more » « less
-
null (Ed.)Control of adaptive walking requires the integration of sensory signals of muscle force and load. We have studied how mechanoreceptors (tibial campaniform sensilla) encode 'naturalistic' stimuli derived from joint torques of stick insects walking on a horizontal substrate. Previous studies showed that forces applied to the legs using the mean torque profiles of a proximal joint were highly effective in eliciting motor activities. However, substantial variations in torque direction and magnitude occurred at the more distal femoro-tibial joint, which can generate braking or propulsive forces and provide lateral stability. To determine how these forces are encoded, we utilized torque waveforms of individual steps that had maximum values in stance in the directions of flexion or extension. Analysis of kinematic data showed that the torques in different directions tended to occur in different ranges of joint angles. Variations within stance were not accompanied by comparable changes in joint angle but often reflected vertical ground reaction forces and leg support of body load. Application of torque waveforms elicited sensory discharges with variations in firing frequency similar to those seen in freely walking insects. All sensilla directionally encoded the dynamics of force increases and showed hysteresis to transient force decreases. Smaller receptors exhibited more tonic firing. Our findings suggest that dynamic sensitivity in force feedback can modulate ongoing muscle activities to stabilize distal joints when large forces are generated at proximal joints. Further, use of 'naturalistic' stimuli can reproduce characteristics seen in freely moving animals that are absent in conventional restrained preparations.more » « less
