Sensorimotor System Research Articles

FNIRS-based adaptive visuomotor task improves sensorimotor cortical activation

Objective. Investigating how to promote the functional activation of the central sensorimotor system is an important goal in the neurorehabilitation research domain. We aim to validate the effectiveness of facilitating cortical excitability using a closed-loop visuomotor task, in which the task difficulty is adaptively adjusted based on an individual’s sensorimotor cortical activation. Approach. We developed a novel visuomotor task, in which subjects moved a handle of a haptic device along a specific path while exerting a constant force against a virtual surface under visual feedback. The difficulty levels of the task were adapted with the aim of increasing the activation of sensorimotor areas, measured non-invasively by functional near-infrared spectroscopy. The changes in brain activation of the bilateral prefrontal cortex, sensorimotor cortex, and the occipital cortex obtained during the adaptive visuomotor task (adaptive group), were compared to the brain activation pattern elicited by the same duration of task with random difficulties in a control group. Main results. During one intervention session, the adaptive group showed significantly increased activation in the bilateral sensorimotor cortex, also enhanced effective connectivity between the prefrontal and sensorimotor areas compared to the control group. Significance. Our findings demonstrated that the functional near-infrared spectroscopy-based adaptive visuomotor task with high ecological validity can facilitate the neural activity in sensorimotor areas and thus has the potential to improve hand motor functions.

Beyond task-space exploration: On the role of variance for motor control and learning.

This conceptual analysis on the role of variance for motor control and learning should be taken as a call to: (a) overcome the classic motor-action controversy by identifying converging lines and mutual synergies in the explanation of motor behavior phenomena, and (b) design more empirical research on low-level operational aspects of motor behavior rather than on high-level theoretical terms. Throughout the paper, claim (a) is exemplified by deploying the well-accepted task-space landscape metaphor. This approach provides an illustration not only of a dynamical sensorimotor system but also of a structure of internal forward models, as they are used in more cognitively rooted frameworks such as the theory of optimal feedback control. Claim (b) is put into practice by, mainly theoretically, substantiating a number of predictions for the role of variance in motor control and learning that can be derived from a convergent perspective. From this standpoint, it becomes obvious that variance is neither generally “good” nor generally “bad” for sensorimotor learning. Rather, the predictions derived suggest that specific forms of variance cause specific changes on permanent performance. In this endeavor, Newell’s concept of task-space exploration is identified as a fundamental learning mechanism. Beyond, we highlight further predictions regarding the optimal use of variance for learning from a converging view. These predictions regard, on the one hand, additional learning mechanisms based on the task-space landscape metaphor—namely task-space formation, task-space differentiation and task-space (de-)composition—and, on the other hand, mechanisms of meta-learning that refer to handling noise as well as learning-to-learn and learning-to-adapt. Due to the character of a conceptual-analysis paper, we grant ourselves the right to be highly speculative on some issues. Thus, we would like readers to see our call mainly as an effort to stimulate both a meta-theoretical discussion on chances for convergence between classically separated lines of thought and, on an empirical level, future research on the role of variance in motor control and learning.

Language and Intelligence: A Relationship Supporting the Embodied Cognition Hypothesis.

Cognitive science has gathered robust evidence supporting the hypothesis that cognitive processes do not occur in an amodal format but take shape through the activation of the sensorimotor systems of the agent body, which works as simulation system upon which concepts, words, and thought are based. However, studies that have investigated the relationship between language and cognitive processes, as both embedded processes, are very rare. In this study, we investigated the hypothesis that intelligence is associated with referential competence, conceived as the ability to find words to refer to our subjective and perceptual experience, and to evoke understanding of this experience in the listener. We administered the WAIS-IV test to 32 nonclinical subjects and collected autobiographical narratives from them through the Relationship Anecdotes Paradigm Interview. The narratives were analyzed linguistically by applying computerized measures of referential competence. Intelligence scores were found to correlate with the use in narratives of words related to somatic and sensory sensations, while they were not associated with other measures of referential competence related to more abstract domains of experience or based on vivid or reflective dimensions of language style. The results support the hypothesis that sensorimotor schemas have an intrinsic role in language and cognition.

Study of reactions of the sensorimotor system in adolescents during and after surgical correction of spinal deformity

BACKGROUND: Little attention has been paid to the study of delayed sensory and motor reactions in adolescents with spinal deformities after surgical treatment.
 AIM: To study the reactions of the sensorimotor system of adolescents after surgical correction of spinal deformity.
 MATERIALS AND METHODS: The state of the sensory and motor spheres was analyzed in the immediate postoperative period in 21 adolescents with idiopathic scoliosis and in 13 with congenital deformities of the spine. A complex of methods involving global and stimulation electroneuromyography was used. The amplitude of motor, reflex potentials and interference electromyogram was evaluated at the maximum arbitrary tension of the lower limb muscles. Using an esthesiometer, thermal pain sensitivity in Th1S2 dermatomes was explored. In the process of surgical correction, intraoperative neuromonitoring was performed with registration of motor evoked potentials of the lower limb muscles.
 RESULTS: At the beginning of surgical intervention, high-amplitude, well-reproducible motor evoked potentials were obtained in all patients. In the group of patients with idiopathic scoliosis, compared with those with congenital deformities, smooth flow of surgery prevailed (p 0.05) without significant changes in motor potentials relative to the baseline (p 0.05). The number of observations of motor potentials decreased in the both groups and did not exceed 10%; the differences were not significant (p 0.05). The study of the reactions of the sensorimotor system in the immediate postoperative period triggered an increase in the amplitude of M-responses of m. rectus femoris, m. flexor digitorum brevis, m. gastrocnemius, and a decrease in the amplitude of the total EMG of m. rectus femoris. Values of H-reflexes remained at the preoperative level. The analysis of thermal pain sensitivity demonstrated the presence of a more pronounced reaction than that of the motor component. Changes in indicators of this type of sensitivity in groups of adolescents with idiopathic and congenital scoliosis were opposite. In idiopathic scoliosis, negative dynamics of the values prevailed, while in adolescents with congenital deformities of the spine, positive dynamics prevailed. This was because the amount of correction of the main and compensatory curves of the deformity in the group with idiopathic scoliosis was 48% greater (p = 0.0004) and 51% greater (p = 0.011), respectively.
 CONCLUSIONS: After surgical correction of spinal deformities in adolescents, the reactions of the sensory system of thermal pain sensitivity were more pronounced than those of the motor sphere.

Stability of inverted pendulum reveals transition between predictive control and impedance control in grip force modulation.

During object manipulation, our sensorimotor sys-tem needs to represent the objects dynamics in order to better control it. This is especially important in the case of grip force control where small forces can cause the object to slip from our fingers, and excessive forces can cause fatigue or even damage the object. While the tradeoff between these two constraints is clear for stable objects, such as lifting a soda can, it is less clear how the sensorimotor system adjusts the grip force for unstable objects. For this purpose, we measured the change in the grip force of individual human participants while they stabilize five different lengths of an inverted pendulum. These lengths set different dynamics of the pendulum, ranging in their degree of controllability. We observed two main states during such manipulation, a marginally stable state of the pendulum and a stabilization state in which participants acted to stabilize the system. While during the stabilization state participants increased their applied grip force, for the stable state we observed a mixed behaviour. For small and less controllable pendulums, grip force increased while for larger pendulums, participants could modulate the the grip force according to the anticipated load forces. Based on these results, we suggest that the pendulum dynamics change the control strategy between predictive control and impedance control.

Quantification of cortical proprioceptive processing through a wireless and miniaturized EEG amplifier.

Corticokinematic coherence (CKC) is computed between limb kinematics and cortical activity (e.g. MEG, EEG), and it can be used to detect, quantify and localize the cortical processing of proprioceptive afference arising from the body. EEG-based studies on CKC have been limited to lab environments due to bulky, non-portable instrumentations. We recently proposed a wireless and miniaturized EEG acquisition system aimed at enabling EEG studies outside the laboratory. The purpose of this work is to compare the EEG-based CKC values obtained with this device with a conventional wired-EEG acquisition system to validate its use in the quantification of cortical proprioceptive processing. Eleven healthy right-handed participants were recruited (six males, four females, age range: 24-40 yr). A pneumatic-movement actuator was used to evoke right index-finger flexion-extension movement at 3 Hz for 4 min. The task was repeated both with the wireless-EEG and wired-EEG devices using the same 30-channel EEG cap preparation. CKC was computed between the EEG and finger acceleration. CKC peaked at the movement frequency and its harmonics, being statistically significant (p < 0.05) in 8-10 out of 11 participants. No statistically significant differences (p < 0.05) were found in CKC strength between wireless-EEG (range 0.03-0.22) and wired-EEG (0.02-0.33) systems, that showed a good agreement between the recording systems (3 Hz: r = 0.57, p = 0.071, 6 Hz: r = 0.82, p = 0.003). As expected, CKC peaked in sensors above the left primary sensorimotor cortex contralateral to the moved right index finger. As the wired-EEG device, the tested wireless-EEG system has proven feasible to quantify CKC, and thus can be used as a tool to study proprioception in the human neocortex. Thanks to its portability, the wireless-EEG used in this study has the potential to enable the examination of cortical proprioception in more naturalistic conditions outside the laboratory environment. Clinical Relevance-Our study will contribute to provide innovative technological foundations for future unobtrusive EEG recordings in naturalistic conditions to examine human sensorimotor system.

Dynamical Synergies in Multidigit Hand Prehension.

Hand prehension requires a highly coordinated control of contact forces. The high dimensional sensorimotor system of the human hand although operates at ease, poses several challenges when replicated for prosthetic control. This study investigates how the dynamical synergies, coordinated spatial patterns of contact forces, contribute to the contact forces in a grasp, and whether the dynamical synergies could potentially serve as candidates for feedforward and feedback mechanisms. Ten right-handed subjects were recruited to grasp and hold mass-varied objects. The contact forces during this multidigit prehension were recorded using an instrumented grip glove. The dynamical synergies were derived using principal component analysis (PCA). The contact force patterns during the grasps were reconstructed using the first few synergies. The significance of the dynamical synergies and the current challenges and possible applications of the dynamical synergies were discussed along with the integration of the dynamical synergies into prosthetics and exoskeletons that can possibly enable near-natural control. This research presents dynamical synergies observed in contact forces during hand grasps. These dynamical synergies could help in improving feedforward force control and sensory feedback in hand prosthetics and exoskeletons.

Learning of Dexterous Object Manipulation in a Virtual Reality Environment.

Humans have unrivalled abilities to perform dexterous object manipulation. This requires the sensorimotor system to quickly adapt to environmental changes and predictively counter act the external disturbances. Many studies have focused on the anticipatory control of digits with real-world experiments. However, examining manipulation using virtual reality with haptic devices expands the possibilities of investigation. In this work, participants grasped and lifted an inverted T-shaped object in a virtual reality setup. The graspable surface of the object was either constrained to a small area or unconstrained. The position of the object's center of mass changed between blocks, and the participants were asked to minimize the rotation of the object during the lift. Our results show that, consistent with the results of real-world experiments, participants gradually learn to adjust the digit positions and forces to predictively compensate for the torque due to the shifted center of mass prior to liftoff. The only major difference found was that the length of trials needed during the adaptation phase to each condition increased from 3 in real-world to 5 in virtual environment.

Sensorimotor development of male and female rats subjected to neonatal anoxia.

Neonatal hypoxic-ischemic encephalopathy (HIE) is one of the most important reasons for morbidity and mortality in term-born infants. HIE impacts early somatic, neurological, and motor development including social. To illustrate the damages in the sensorimotor system, an adapted and validated model of neonatal anoxia is used. This study evaluated the sex differences in Wistar rats, neurological reflex, and motor development at the suckling period. Short- and long-term impairments associated with sex differences were observed. In general, anoxic males were more affected in comparison to their control group and to anoxic females. Long-lasting effects of the injury in adolescent rats predominately affected males. Similar to previous studies, we also found a decrease in the number of the substantia nigra cells in both sexes, compared to their control. So far, the results indicate that HIE caused neurobehavioral alterations and asymmetrical motor behavior with brain damage, possibly related to cognitive impairments previously observed at adolescence. These alterations may represent a useful endpoint for studying the efficacy of potential strategies that may improve the developmental consequences of a perinatal asphyxia insult in humans.

Strike or ball? Batters know it better: an fMRI study of action anticipation in baseball players.

To assess whether the brain processes of action anticipation are modulated differently by perceptual and motor experiences, baseball batters, pitchers, and non-players were asked to predict the fate of pitching actions (strike or ball) while undergoing functional magnetic resonance imaging. Results showed both batters (perceptual experts of pitching action) and pitchers (motor experts) were more accurate than non-players. Furthermore, batters demonstrated higher perceptual sensitivity in discriminating strikes than non-players. All groups engaged the action observation network, putamen, and cerebellum during anticipation, while pitchers showed higher activity than non-players in the left premotor cortex, which has been implicated in the internal simulation of observed action. Only batters exhibited differences in strike versus ball pitches in their left ventral extrastriate cortex, which might be associated with the processing of relevant visual information conveyed by the observed pitcher's movement kinematics and pitch trajectory. Moreover, all groups showed higher activity selectively in the striatum, thalamus, sensorimotor cortices, and cerebellum during correct predictions than during incorrect ones, with most widespread activation in batters, reinforcing the greater involvement of the sensorimotor system in perceptual experience. Our findings demonstrate that perceptual experience might enhance action anticipation ability to a greater extent than motor experience, with overlapping but specific neural underpinnings.

A Cross-Cultural Replication on Humanness Attribution

Abstract. Previous research showed that targets achieving (vs. not) a goal were ascribed more humanness. We conceptually replicate previous studies by involving a population of English and Arabic speakers to test cross-cultural replicability of the effect and the contribution of sensorimotor systems in agency representation. Participants ( Ntot = 637) saw animations, where goal achievement and trajectory were manipulated. They evaluated agency, communion, humanness, and attitudes (respect and liking) toward presented targets. Goal achievement versus failure but not movement trajectory increased agency and communion ratings, which in turn affected humanness, respect, and liking (Study 1 and Study 2). Goal manipulation also directly affected humanness ratings (Study 2). Altogether, our findings suggest a superior role of success over trajectory manipulation in perceiving inanimate objects as having humanness.

Misencoding of ankle joint angle control system via cutaneous afferents reflex pathway in chronic ankle instability.

This study aimed to investigate how the cutaneous reflexes in the peroneus longus (PL) muscle are affected by changing the ankle joint position in patients with chronic ankle instability (CAI). We also investigated the correlation between the degree of reflex modulation and angle position sense of the ankle joint. The participants were 19 patients with CAI and 20 age-matched controls. Cutaneous reflexes were elicited by applying non-noxious electrical stimulation to the sural nerve at the ankle joint in the neutral standing and eversion/inversion standing positions. The suppressive middle latency cutaneous reflex (MLR; ~ 70-120ms) and angle position sense of the ankle joint were assessed. During neutral standing, the gain of the suppressive MLR was more prominent in the CAI patients than in controls, although no significant difference was seen during 30° inversion standing. In addition, the ratios of the suppressive MLR and background electromyography in a neutral position were significantly larger than those at the 15°, 25°, and 30° inversion positions in CAI patients. No such difference was seen in control individuals. Furthermore, the correlations between reflex modulation degree and position sense error were quite different in CAI patients compared to controls. These findings suggest that the sensory-motor system was deteriorated in CAI patients due to changes in the PL cutaneous reflex pathway excitability and position sense of the ankle joint.

What does it actually mean that Premotor Theory is about embodied attention?

ABSTRACT One of the most vigorously debated issues in attention labs concerns the nature of the coupling between the sensory-motor system and covert spatial attention. Proponents of the Premotor Theory of Attention (PToA) claim that attention should be accounted for in terms of motor preparation for goal-directed actions such as eye or hand movements. For others, it is a supramodal psychological entity that is independent of our sensorimotor machinery. Both parties also seek to articulate this controversy in terms of cognitive science as a question of whether attention is embodied or disembodied cognition. The PToA is thought to advocate the embodiment of attention. We, however, are inclined to think that a more nuanced view of this conceptual framework is needed to yield some insights into how radical it actually is in terms of embodied cognition (EC). We argue that PToA still seems to bear some features of weak embodiment. Yet we suggest that it has the potential to build a stronger concept of attention as EC, but somewhat paradoxically this would require giving attention deflationary treatment and reconstructing it in other terms. We propose that the notion of pre-reflective intentionality might serve as an inspiration to this end.

Uneven Treadmill Training for Rehabilitation of Lateral Ankle Sprains and Chronic Ankle Instability: Protocol for a Pragmatic Randomized Controlled Trial.

BackgroundLateral ankle sprains (LASs) are common injuries among military service members. Approximately 40% of individuals with an LAS progress to develop chronic ankle instability (CAI), a condition that results in substantial mechanical and neurophysiological impairment and activity limitation. Since proprioceptive and balance training improve functional outcomes and prevent secondary injury following LAS, they are recommended in clinical practice. Uneven treadmills are an innovative modality that challenge the sensorimotor system while performing an ecologically valid task simulating environments frequently encountered by service members with LAS and CAI.ObjectiveThe aim of this study is to evaluate whether the inclusion of uneven treadmill training in standard rehabilitation can improve clinical, functional, biomechanical, and patient-reported outcomes compared with the standard of care alone in service members with LAS and CAI. The prophylactic effects of treatment on secondary injury and identification of any contributing or mediating factors that influence outcomes following treatment will also be evaluated. We hypothesize that service members receiving uneven treadmill training will demonstrate greater improvements in clinical and instrumented measures of impairment, patient-reported function, and lower risk of injury recurrence than the control group immediately post and 18 months following treatment.MethodsA multisite, parallel randomized clinical trial will be performed among service members aged 18-49 years being treated for LAS and CAI in military treatment facilities in the United States. Participants randomly assigned and allocated to receive the experimental intervention will be provided up to 12 sessions of training on an uneven terrain treadmill over a 6-week treatment course to supplement standard rehabilitation care. Treatment intensity of the rehabilitation exercises and treadmill training will be progressed on the basis of patient-perceived intensity and treatment responses. Outcome measures will include patient-reported outcomes, functional assessments, performance measures, and biomechanical measures. Investigators collecting outcome measures will be blinded to treatment allocation. Reinjury rates and patient-reported outcomes of function will be tracked over 18 months following treatment.ResultsThe project was funded in September 2020. Patient recruitment began in November 2021, with 3 participants enrolled as of February 2022. Dissemination of the main study findings is anticipated in 2024.ConclusionsThis study will assess the impact of an innovative uneven-terrain treadmill on treatment outcomes in the rehabilitation of service members with LAS and CAI. The results of this study will be used to inform rehabilitation practices and to potentially improve functional outcomes and secondary prevention in this patient population.Trial RegistrationClinicalTrials.gov NCT04999904; https://clinicaltrials.gov/ct2/show/NCT04999904?term=NCT04999904International Registered Report Identifier (IRRID)DERR1-10.2196/38442

Influences of hand action on the processing of symbolic numbers: A special role of pointing?

Embodied and grounded cognition theories state that cognitive processing is built upon sensorimotor systems. In the context of numerical cognition, support to this framework comes from the interactions between numerical processing and the hand actions of reaching and grasping documented in skilled adults. Accordingly, mechanisms for the processing of object size and location during reach and grasp actions might scaffold the development of mental representations of numerical magnitude. The present study exploited motor adaptation to test the hypothesis of a functional overlap between neurocognitive mechanisms of hand action and numerical processing. Participants performed repetitive grasping of an object, repetitive pointing, repetitive tapping, or passive viewing. Subsequently, they performed a symbolic number comparison task. Importantly, hand action and number comparison were functionally and temporally dissociated, thereby minimizing context-based effects. Results showed that executing the action of pointing slowed down the responses in number comparison. Moreover, the typical distance effect (faster responses for numbers far from the reference as compared to close ones) was not observed for small numbers after pointing, while it was enhanced by grasping. These findings confirm the functional link between hand action and numerical processing, and suggest new hypotheses on the role of pointing as a meaningful gesture in the development and embodiment of numerical skills.

Philosophical Specification of Empathetic Ethical Artificial Intelligence

In order to construct an ethical artificial intelligence (AI) two complex problems must be overcome. Firstly, humans do not consistently agree on what is or is not ethical. Second, contemporary AI and machine learning methods tend to be blunt instruments which either search for solutions within the bounds of predefined rules, or mimic behaviour. An ethical AI must be capable of inferring unspoken rules, interpreting nuance and context, possess and be able to infer intent, and explain not just its actions but its intent. Using enactivism, semiotics, perceptual symbol systems and symbol emergence, we specify an agent that learns not just arbitrary relations between signs but their meaning in terms of the perceptual states of its sensorimotor system. Subsequently it can learn what is meant by a sentence and infer the intent of others in terms of its own experiences. It has malleable intent because the meaning of symbols changes as it learns, and its intent is represented symbolically as a goal. As such it may learn a concept of what is most likely to be considered ethical by the majority within a population of humans, which may then be used as a goal. The meaning of abstract symbols is expressed using perceptual symbols of raw sensorimotor stimuli as the weakest (consistent with Ockham's Razor) necessary and sufficient concept, an intensional definition learned from an ostensive definition, from which the extensional definition or category of all ethical decisions may be obtained. Because these abstract symbols are the same for both situation and response, the same symbol is used when either performing or observing an action. This is akin to mirror neurons in the human brain. Mirror symbols may allow the agent to empathise, because its own experiences are associated with the symbol, which is also associated with the observation of another agent experiencing something that symbol represents.

Separating Stimulus-Induced and Background Components of Dynamic Functional Connectivity in Naturalistic fMRI.

We consider the challenges in extracting stimulus-related neural dynamics from other intrinsic processes and noise in naturalistic functional magnetic resonance imaging (fMRI). Most studies rely on inter-subject correlations (ISC) of low-level regional activity and neglect varying responses in individuals. We propose a novel, data-driven approach based on low-rank plus sparse ( [Formula: see text]) decomposition to isolate stimulus-driven dynamic changes in brain functional connectivity (FC) from the background noise, by exploiting shared network structure among subjects receiving the same naturalistic stimuli. The time-resolved multi-subject FC matrices are modeled as a sum of a low-rank component of correlated FC patterns across subjects, and a sparse component of subject-specific, idiosyncratic background activities. To recover the shared low-rank subspace, we introduce a fused version of principal component pursuit (PCP) by adding a fusion-type penalty on the differences between the columns of the low-rank matrix. The method improves the detection of stimulus-induced group-level homogeneity in the FC profile while capturing inter-subject variability. We develop an efficient algorithm via a linearized alternating direction method of multipliers to solve the fused-PCP. Simulations show accurate recovery by the fused-PCP even when a large fraction of FC edges are severely corrupted. When applied to natural fMRI data, our method reveals FC changes that were time-locked to auditory processing during movie watching, with dynamic engagement of sensorimotor systems for speech-in-noise. It also provides a better mapping to auditory content in the movie than ISC.

A review of upper limb robot assisted therapy techniques and virtual reality applications

Impairments in the sensorimotor system negatively impact the ability of individuals to perform daily activities autonomously. Upper limb rehabilitation for stroke survivors and cerebral palsy (CP) children is essential to enhance independence and quality of life. Robot assisted therapy has been a bright solution in the last two decades to promote the recovery process for neurological disorders patients. Nevertheless, defining the optimum intervention of robot assisted therapy (RAT) in different cases is not clear yet. With this aim, the presented study reviewed the current literature on RAT protocols for upper limb impairments and the effects of RAT on recovery outcomes. A literature search was conducted using different search engines, reviews, and studies. This study presents an overview of fourteen robotic devices used in the rehabilitation field and seventeen clinical trials using commercially available devices during the last three years. A discussion about reaching an efficient rehabilitation process based on different aspects such as clinical setting and training modes has been introduced. This review identifies the limitations of RAT to lay the foundation for more effective neuromotor disorders rehabilitation. Finally, using virtual reality (VR) as an assisting feature in RAT improves the whole process of recovering motor functionality.

Dynamical Synergies of Multidigit Hand Prehension.

Hand prehension requires highly coordinated control of contact forces. The high-dimensional sensorimotor system of the human hand operates at ease, but poses several challenges when replicated in artificial hands. This paper investigates how the dynamical synergies, coordinated spatiotemporal patterns of contact forces, contribute to the hand grasp, and whether they could potentially capture the force primitives in a low-dimensional space. Ten right-handed subjects were recruited to grasp and hold mass-varied objects. The contact forces during this multidigit prehension were recorded using an instrumented grip glove. The dynamical synergies were derived using principal component analysis (PCA). The contact force patterns during the grasps were reconstructed using the first few synergies. The significance of the dynamical synergies, the influence of load forces and task configurations on the synergies were explained. This study also discussed the contribution of biomechanical constraints on the first few synergies and the current challenges and possible applications of the dynamical synergies in the design and control of exoskeletons. The integration of the dynamical synergies into exoskeletons will be realized in the near future.

Kinematic trajectories in response to speed perturbations in walking suggest modular task-level control of leg angle and length.

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.