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Perceptual decision-making involves multiple cognitive processes, including accumulation of sensory evidence, planning, and executing a motor action. How these processes are intertwined is unclear; some models assume that decision-related processes precede motor execution, whereas others propose that movements reflecting ongoing decision processes occur before commitment to a choice. Here we combine two complementary methods to study the relationship between decision processes and the movements leading up to a choice. The first is a free-response pulse-based evidence accumulation task, in which stimuli continue until choice is reported, and the second is a motion-based drift diffusion model (mDDM), in which movement variables from video pose estimation constrain decision parameters on a trial-by-trial basis. We find that the mDDM provides a better fit to rats' decisions in the free-response accumulation task than traditional drift diffusion models. Interestingly, on each trial we observed a period, before choice, that was characterized by head immobility. The length of this period was positively correlated with the rats' decision bounds, and stimuli presented during this period had the greatest impact on choice. Together these results support a model in which internal decision dynamics are reflected in movements and demonstrate that inclusion of movement parameters improves the performance of diffusion-to-bound decision models. NEW & NOTEWORTHY In this study we combine a novel pulse-based evidence accumulation task with a newly developed motion-based drift diffusion model (mDDM). In this model, we incorporate movement parameters derived from high-resolution video data to estimate parameters of the model on a trial-by-trial basis. We find that this new model is an improved description of animal choice behavior.

Many recent studies indicate that control of decisions and actions is integrated during interactive behavior. Among these, several carried out in humans and monkeys conclude that there is a coregulation of choices and movements. Another perspective, based on human data only, proposes a decoupled control of decision duration and movement speed, allowing, for instance, trading decision duration for movement duration when time pressure increases. Crucially, it is not currently known whether this ability to flexibly dissociate decision duration from movement speed is specific to humans, whether it can vary depending on the context in which a task is performed, and whether it is stable over time. These are important questions to address, especially to rely on monkey electrophysiology to infer the neural mechanisms of decision-action coordination in humans. To do so, we trained two macaque monkeys in a perceptual decision-making task and analyzed data collected over multiple behavioral sessions. Our findings reveal a strong and complex relationship between decision duration and movement vigor. Decision duration and action duration can covary but also "compensate" each other. Such integrated but decoupled control of decisions and actions aligns with recent studies in humans, validating the monkey model in electrophysiology as a means of inferring neural mechanisms in humans. Crucially, we demonstrate for the first time that this control can evolve with experience, in an adapted manner. Together, the present findings contribute to deepening our understanding of the integrated control of decisions and actions during interactive behavior. NEW & NOTEWORTHY The mechanism by which the integrated control of decisions and actions occurs, coupled or interactive but decoupled, is debated. In the present study, we show in monkeys that decisions and actions influence each other in a decoupled way. For the first time, we also demonstrate that this control can evolve depending the subject's experience, allowing the trade of movement time for decision time and limiting the temporal discounting of reward value.

Holding still and aiming reaches to spatial targets may depend on distinct neural circuits. Using automated homecage training and a sensitive joystick, we trained freely moving mice to contact a joystick, hold their forelimb still, and then reach to rewarded target locations. Mice learned the task by initiating forelimb sequences with clearly resolved submillimeter-scale micromovements followed by millimeter-scale reaches to learned spatial targets. Hundreds of thousands of trajectories were decomposed into millions of kinematic submovements, while photoinhibition was used to test roles of motor cortical areas. Inactivation of both caudal and rostral forelimb areas preserved the ability to produce aimed reaches, but reduced reach speed. Inactivation specifically of contralateral caudal forelimb area (CFA) additionally impaired the ability to aim corrective submovements to remembered locations following target undershoots. Our findings show that motor cortical inactivations reduce the gain of forelimb movements but that inactivation specifically of contralateral CFA impairs corrective movements important for reaching a target location. NEW & NOTEWORTHY To test the role of different cortical areas in holding still and reaching to targets, this study combined home-cage training with optogenetic silencing as mice engaged in a learned center-out-reach task. Inactivation specifically of contralateral caudal forelimb area (CFA) impaired corrective movements necessary to reach spatial targets to earn reward.

Information about another person's movement kinematics obtained through visual observation activates brain regions involved in motor learning. Observation-related changes in these brain areas are associated with adaptive changes to feedforward neural control of muscle activation and behavioral improvements in limb movement control. However, little is known about the stability of these observation-related effects over time. Here, we used force channel trials to probe changes in lateral force production at various time points (1 min, 10 min, 30 min, 60 min, 24 h) after participants either physically performed, or observed another individual performing upper limb reaching movements that were perturbed by novel, robot-generated forces (a velocity-dependent force-field). Observers learned to predictively generate directionally and temporally specific compensatory forces during reaching, consistent with the idea that they acquired an internal representation of the novel dynamics. Participants who physically practiced in the force-field showed adaptation that was detectable at all time points, with some decay detected after 24 h. Observation-related adaptation was less temporally stable in comparison, decaying slightly after 1 h and undetectable at 24 h. Observation induced less adaptation overall than physical practice, which could explain differences in temporal stability. Visually acquired representations of movement dynamics are retained and continue to influence behavior for at least 1 h after observation. NEW & NOTEWORTHY We used force channel probes in an upper limb force-field reaching task in humans to compare the durability of learning-related changes that occurred through visual observation to those after physical movement practice. Visually acquired representations of movement dynamics continued to influence behavior for at least 1 h after observation. Our findings point to a 1-h window during which visual observation of another person could play a role in motor learning.

In 1996, we published a review article (Marder E, Calabrese RL. Physiol Rev 76: 687-717, 1996) describing the state of knowledge about the structure and function of the central pattern-generating circuits important for producing rhythmic behaviors. Although many of the core questions persist, much has changed since 1996. Here, we focus on newer studies that reveal ambiguities that complicate understanding circuit dynamics, despite the enormous technical advances of the recent past. In particular, we highlight recent studies of animal-to-animal variability and our understanding that circuit rhythmicity may be supported by multiple state-dependent mechanisms within the same animal and that robustness and resilience in the face of perturbation may depend critically on the presence of modulators and degenerate circuit mechanisms. Additionally, we highlight the use of computational models to ask whether there are generalizable principles about circuit motifs that can be found across rhythmic motor systems in different animal species.

Localizing one's body parts is important for movement control and motor learning. Recent studies have shown that the precision with which people localize their hand places constraints on motor adaptation. Although these studies have assumed that hand localization remains equally precise across learning, we show that precision decreases rapidly during early motor learning. In three experiments, healthy young participants ( n = 92) repeatedly adapted to a 45° visuomotor rotation for a cycle of two to four reaches, followed by a cycle of two to four reaches with veridical feedback. Participants either used an aiming strategy that fully compensated for the rotation ( experiment 1 ), or always aimed directly at the target, so that adaptation was implicit ( experiment 2 ). We omitted visual feedback for the last reach of each cycle, after which participants localized their unseen hand. We observed an increase in the variability of angular localization errors when subjects used a strategy to counter the visuomotor rotation ( experiment 1 ). This decrease in precision was less pronounced in the absence of reaiming ( experiment 2 ), and when subjects knew that they would have to localize their hand on the upcoming trial, and could thus focus on hand position ( experiment 3 ). We propose that strategic reaiming decreases the precision of perceived hand position, possibly due to attention to vision rather than proprioception. We discuss how these dynamics in precision during early motor learning could impact on motor control and shape the interplay between implicit and strategy-based motor adaptation. NEW & NOTEWORTHY Recent studies indicate that the precision with which people localize their hand limits implicit visuomotor learning. We found that localization precision is not static, but decreases early during learning. This decrease is pronounced when people apply a reaiming strategy to compensate for a visuomotor perturbation and is partly resistant to allocation of attention to the hand. We propose that these dynamics in position sense during learning may influence how implicit and strategy-based motor adaption interact.

The flexibility of the motor system to adjust a planned action before or during the execution of the movement in response to sensory information is critical for preventing errors in motor control. As individuals age, this function declines, leading to an increased incidence of motor errors. Although sensory processing and cognitive decline are known contributors to this impairment, here, we test the hypothesis that repetition of context-specific planned actions interferes with the adjustment of feedforward motor commands. Younger and older participants were instructed to grasp and lift a T-shaped object with a concealed, off-sided center of mass and minimize its roll through anticipatory force control, relying predominantly on predictive model-driven planning (i.e., sensorimotor memories) developed through repeated lifts. We selectively manipulate the number of trial repeats with the center of mass on one side before switching it to the other side of the T-shaped object. The results showed that increasing the number of repetitions improved performance in manipulating an object with a given center of mass but led to increased errors when the object's center of mass was switched. This deleterious effect of repetition on feedforward motor adjustment was observed in younger and older adults. Critically, we show these effects on an internal model-driven motor planning task that relies predominantly on sensorimotor memory, with no differences in sensory inputs from the repetition manipulation. The findings indicate that feedforward motor adjustments are hampered by repetitive stereotyped planning and execution of motor behavior. NEW & NOTEWORTHY Adjusting planned actions in response to sensory stimuli degrades with age contributing to increased incidence of errors ranging from clumsy spills to catastrophic falls. Multiple factors likely contribute to age-related motor inflexibility, including sensory- and cognition-supporting system declines. Here, we present compelling evidence for repetition to disrupt feedforward adjusting of motor commands in younger and older adults, which suggests increases in stereotypy as a deleterious potentiator of motor control errors.

The aim of this study was to evaluate mirror visual feedback (MVF) as a training tool for brain-computer interface (BCI) users. This is because approximately 20-30% of subjects require more training to operate a BCI system using motor imagery. Electroencephalograms (EEGs) were recorded from 18 healthy subjects, using event-related desynchronization (ERD) to observe the responses during the movement or movement intention of the hand for the conditions of control, imagination, and the MVF with the mirror box. We constituted two groups: group 1 : control, imagination, and MVF; group 2 : control, MVF, and imagination. There were significant differences in imagination conditions between groups using MVF before or after imagination (right-hand, P = 0.0403; left-hand, P = 0.00939). The illusion of movement through MVF is not possible in all subjects, but even in those cases, we found an increase in imagination when the subject used the MVF previously. The increase in the r 2 s of imagination in the right and left hands suggests cross-learning. The increase in motor imagery recorded with EEG after MVF suggests that the mirror box made it easier to imagine movements. Our results provide evidence that the MVF could be used as a training tool to improve motor imagery. NEW & NOTEWORTHY The increase in motor imagery recorded with EEG after MVF (mirror visual feedback) suggests that the mirror box made it easier to imagine movements. Our results demonstrate that MVF could be used as a training tool to improve motor imagery.

Motor disturbances predominantly characterize hypoxic-ischemic encephalopathy (HIE). Among its intervention methods, environmental enrichment (EE) is strictly considered a form of sensory intervention. However, limited research uses EE as a single sensory input intervention to validate outcomes postintervention. A Sprague-Dawley rat model subjected to left common carotid artery ligation and exposure to oxygen-hypoxic conditions is used in this study. EE was achieved by enhancing the recreational and stress-relief items within the cage, increasing the duration of sunlight, colorful items exposure, and introducing background music. JZL184 (JZL) was administered as neuroprotective drugs. EE was performed 21 days postoperatively and the rats were randomly assigned to the standard environment and EE groups, the two groups were redivided into control, JZL, and vehicle injection subgroups. The Western blotting and behavior test indicated that EE and JZL injections were efficacious in promoting cognitive function in rats following HIE. In addition, the motor function performance in the EE-alone intervention group and the JZL-alone group after HIE was significantly improved compared with the control group. The combined EE and JZL intervention group exhibited even more pronounced improvements in these performances. EE may enhance motor function through sensory input different from the direct neuroprotective effect of pharmacological treatment. NEW & NOTEWORTHY Rarely does literature assess motor function, even though it is common after hypoxia ischemic encephalopathy (HIE). Previously used environmental enrichment (EE) components have not been solely used as sensory inputs. Physical factors were minimized in our study to observe the effects of purely sensory inputs.

Regular exercise has numerous health benefits, but the human population displays significant variability in exercise participation. Rodent models, such as voluntary wheel running (VWR) in rats, can provide insight into the underlying mechanisms of exercise behavior and its regulation. In this study, we focused on the role of estrogen on VWR in female rats. Female rats run more than males, and we aimed to determine to what extent running levels in females were regulated by estrogen signaling. The running behavior of rats (duration, speed, and total distance run) was measured under normal physiological conditions, ovariectomy (OVX), and estrogen replacement in an OVX background. Results show cyclic variations in running linked to the estrous cycle. Ovariectomy markedly reduced running and eliminated the cyclic pattern. Estrogen replacement through estradiol benzoate (EB) injections and osmotic minipumps reinstated running activity to pre-OVX levels and restored the cyclic pattern. Importantly, individual differences and ranking are preserved such that high versus low runners before OVX remain high and low runners after treatment. Further analysis revealed that individual variation in running distance was primarily caused by rats running different speeds, but rats also varied in running duration. However, it is noteworthy that this model also displays features distinct from estrogen-driven running behavior under physiological conditions, notably a delayed onset and a broader duration of running activity. Collectively, this estrogen causality VWR model presents a unique opportunity to investigate sex-specific mechanisms that control voluntary physical activity. NEW & NOTEWORTHY This study investigates estrogen's role in voluntary wheel running (VWR) behavior in female rats. Female rats exhibit greater running than males, with estrogen signaling regulating this activity. The estrous cycle influences running, whereas ovariectomy reduces it, and estrogen replacement restores it, maintaining individual differences under all conditions. Both running speed and duration contribute to VWR variations. These findings emphasize individual estrogen regulation in female exercise and provide an estrogen replacement animal model for investigating neurobiological underpinnings that drive voluntary exercise behavior.

We present the first demonstration of chronic in vivo imaging of microglia in mice undergoing voluntary wheel running. We find that healthy mice undergoing voluntary wheel running have similar microglia dynamics, morphologies, and responses to injury when compared to sedentary mice. This suggests that exercise over a period of 1 mo does not grossly alter cortical microglial phenotypes and that exercise may exert its beneficial effects on the brain through other mechanisms. Future work examining how microglia dynamics may be altered during exercise in disease or injury models could provide further insights into the therapeutic benefit of exercise. NEW & NOTEWORTHY We demonstrate the first use of chronic in vivo imaging of microglia over time during physical exercise. We found that microglia movement, morphology, and process motility were remarkably stable during voluntary wheel running (VWR). Additionally, microglia in running mice respond similarly to laser ablation injury compared to sedentary mice. These findings indicate that VWR does not induce changes in microglia dynamics in healthy adults. Exercise may elicit positive effects on the brain through other mechanisms.

Cardiovascular disease is an enormous public health problem, particularly in older populations. Exercise is the most potent cardioprotective intervention identified to date, with exercise in the juvenile period potentially imparting greater protection, given the plasticity of the developing heart. To test the hypothesis that voluntary wheel running early in life would be cardioprotective later in life when risk for disease is high, we provided male and female juvenile (3 wk old) mice access to a running wheel for 2 wk. Mice then returned to a home cage to age to adulthood (4-6 mo) before exposure to isoproterenol (ISO) to induce cardiac stress. Cardiac function and remodeling were compared with sedentary control mice, sedentary mice exposed to ISO, and mice that exercised in adulthood immediately before ISO. Early in life activity protected against ISO-induced stress as evidenced by attenuated cardiac mass, myocyte size, and fibrosis compared with sedentary mice exposed to ISO. ISO-induced changes in cardiac function were ameliorated in male mice that engaged in wheel running, with ejection fraction and fractional shortening reversed to control values. Adrenergic receptor expression was downregulated in juvenile male runners. This suppression persisted in adulthood following ISO, providing a putative mechanism by which exercise in the young male heart provides resilience to cardiac stress later in life. Together, we show that activity early in life induces persistent cardiac changes that attenuate ISO-induced stress in adulthood. Identification of the mechanisms by which early in life exercise is protective will yield valuable insights into how exercise is medicine across the life course. NEW & NOTEWORTHY Voluntary wheel running activity early in life induces persistent changes in the heart that attenuate isoproterenol-induced hypertrophy and fibrosis in adulthood. Though the mechanisms of this protection remain incompletely understood, activity-induced downregulation of adrenergic receptor expression early in life may contribute to later protection against adrenergic stress. Together these data suggest that efforts to maintain an active lifestyle early in life may have long-lasting cardioprotective benefits.

We sought to determine the effects of long-term voluntary wheel running on markers of long interspersed nuclear element-1 (L1) in skeletal muscle, liver, and the hippocampus of female rats. In addition, markers of the cGAS-STING DNA-sensing pathway that results in inflammation were interrogated. Female Lewis rats ( n = 34) were separated into one of three groups including a 6-mo-old group to serve as a young comparator group (CTL, n = 10), a group that had access to a running wheel for voluntary wheel running (EX, n = 12), and an age-matched group that did not (SED, n = 12). Both SED and EX groups were carried out from 6 mo to 15 mo of age. There were no significant differences in L1 mRNA expression for any of the tissues between groups. Methylation of the L1 promoter in the soleus and hippocampus was significantly higher in SED and EX than in CTL group ( P < 0.05). ORF1p expression was higher in older SED and EX rats than in CTL rats for every tissue ( P < 0.05). There were no differences between groups for L1 mRNA or cGAS-STING pathway markers. Our results suggest there is an increased ORF1 protein expression across tissues with aging that is not mitigated by voluntary wheel running. In addition, although previous data imply that L1 methylation changes may play a role in acute exercise for L1 RNA expression, this does not seem to occur during extended periods of voluntary wheel running.

Reinforcement learning has been used as an experimental model of motor skill acquisition, where at times movements are successful and thus reinforced. One fundamental problem is to understand how humans select exploration over exploitation during learning. The decision could be influenced by factors such as task demands and reward availability. In this study, we applied a clustering algorithm to examine how a change in the accuracy requirements of a task affected the choice of exploration over exploitation. Participants made reaching movements to an unseen target using a planar robot arm and received reward after each successful movement. For one group of participants, the width of the hidden target decreased after every other training block. For a second group, it remained constant. The clustering algorithm was applied to the kinematic data to characterize motor learning on a trial-to-trial basis as a sequence of movements, each belonging to one of the identified clusters. By the end of learning, movement trajectories across all participants converged primarily to a single cluster with the greatest number of successful trials. Within this analysis framework, we defined exploration and exploitation as types of behavior in which two successive trajectories belong to different or similar clusters, respectively. The frequency of each mode of behavior was evaluated over the course of learning. It was found that by reducing the target width, participants used a greater variety of different clusters and displayed more exploration than exploitation. Excessive exploration relative to exploitation was found to be detrimental to subsequent motor learning. NEW & NOTEWORTHY The choice of exploration versus exploitation is a fundamental problem in learning new motor skills through reinforcement. In this study, we employed a data-driven approach to characterize movements on a trial-by-trial basis with an unsupervised clustering algorithm. Using this technique, we found that changes in task demands and, in particular, in the required accuracy of movements, influenced the ratio of exploration to exploitation. This analysis framework provides an attractive tool to investigate mechanisms of explorative and exploitative behavior while studying motor learning.

Acquisition of multiple motor skills without interference is a remarkable ability in daily life. During adaptation to opposing perturbations, a common paradigm to study this ability, each perturbation can be successfully learned when a contextual follow-through movement is associated with the direction of the perturbation. It is still unclear, however, to what extent this learning engages the cognitive explicit process and the implicit process. Here, we untangled the individual contributions of the explicit and implicit components while participants learned opposing visuomotor perturbations, with a second unperturbed follow-through movement. In experiment 1 , we replicated previous adaptation results and showed that follow-through movements also allow learning for opposing visuomotor rotations. For one group of participants in experiment 2 , we isolated strategic explicit learning, while for another group we isolated the implicit component. Our data showed that opposing perturbations could be fully learned by explicit strategies, but when strategy was restricted, distinct implicit processes contributed to learning. In experiment 3 , we examined whether learning is influenced by the disparity between the follow-through contexts. We found that the location of follow-through targets had little effect on total learning, yet it led to more instances in which participants failed to learn the task. In experiment 4 , we explored the generalization capability to untrained targets. Participants showed near-flat generalization of the implicit and explicit processes. Overall, our results indicate that follow-through contextual cues might activate, in part, top-down cognitive factors that influence not only the dynamics of the explicit learning but also the implicit process. NEW & NOTEWORTHY Acquisition of multiple motor skills is a remarkable ability in everyday activities. Yet, the contribution of implicit and explicit learning processes during learning of multiple motor skills is still unclear. We sought to dissociate this ability during learning opposing visuomotor rotations, each associated with a contextual follow-through movement. We show that follow-through contextual cues influence activity of both implicit and explicit processes, suggesting that context-based top-down cognitive factors influence not only the explicit learning but also the implicit process.

Skilled movements result from a mixture of feedforward and feedback mechanisms conceptualized by internal models. These mechanisms subserve both motor execution and motor imagery. Current research suggests that imagery allows updating feedforward mechanisms, leading to better performance in familiar contexts. Does this still hold in radically new contexts? Here, we test this ability by asking participants to imagine swinging arm movements around shoulder in normal gravity condition and in microgravity in which studies showed that movements slow down. We timed several cycles of actual and imagined arm pendular movements in three groups of subjects during parabolic flight campaign. The first, control, group remained on the ground. The second group was exposed to microgravity but did not imagine movements inflight. The third group was exposed to microgravity and imagined movements inflight. All groups performed and imagined the movements before and after the flight. We predicted that a mere exposure to microgravity would induce changes in imagined movement duration. We found this held true for the group who imagined the movements, suggesting an update of internal representations of gravity. However, we did not find a similar effect in the group exposed to microgravity despite the fact that the participants lived the same gravitational variations as the first group. Overall, these results suggest that motor imagery contributes to update internal representations of the considered movement in unfamiliar environments, while a mere exposure proved to be insufficient. NEW & NOTEWORTHY Gravity strongly affects the way movements are performed. How internal models process this information to adapt behavior to novel contexts is still unknown. The microgravity environment itself does not provide enough information to optimally adjust the period of natural arm swinging movements to microgravity. However, motor imagery of the task while immersed in microgravity was sufficient to update internal models. These results show that actually executing a task is not necessary to update graviception.

A consistent finding in sensorimotor adaptation is a persistent undershoot of full compensation, such that performance asymptotes with residual errors greater than seen at baseline. This behavior has been attributed to limiting factors within the implicit adaptation system, which reaches a suboptimal equilibrium between trial-by-trial learning and forgetting. However, recent research has suggested that allowing longer motor planning periods prior to movement eliminates these residual errors. The additional planning time allows required cognitive processes to be completed before movement onset, thus increasing accuracy. Here, we looked to extend these findings by investigating the relationship between increased motor preparation time and the size of imposed visuomotor rotation (30°, 45°, or 60°), with regard to the final asymptotic level of adaptation. We found that restricting preparation time to 0.35 s impaired adaptation for moderate and larger rotations, resulting in larger residual errors compared to groups with additional preparation time. However, we found that even extended preparation time failed to eliminate persistent errors, regardless of magnitude of cursor rotation. Thus, the asymptote of adaptation was significantly less than the degree of imposed rotation, for all experimental groups. In addition, there was a positive relationship between asymptotic error and implicit retention. These data suggest that a prolonged motor preparation period is insufficient to reliably achieve complete adaptation, and therefore, our results suggest that factors beyond that of planning time contribute to asymptotic adaptation levels. NEW & NOTEWORTHY Residual errors in sensorimotor adaptation are commonly attributed to an equilibrium between trial-by-trial learning and forgetting. Recent research suggested that allowing sufficient time for mental rotation eliminates these errors. In a number of experimental conditions, we show that although restricted motor preparation time does limit adaptation-consistent with mental rotation-extending preparation time fails to eliminate the residual errors in motor adaptation.

Brain systems supporting body movement are active during music listening in the absence of overt movement. This covert motor activity is not well understood, but some theories propose a role in auditory timing prediction facilitated by motor simulation. One question is how music-related covert motor activity relates to motor activity during overt movement. We address this question using scalp electroencephalogram by measuring mu rhythms-cortical field phenomena associated with the somatomotor system that appear over sensorimotor cortex. Lateralized mu enhancement over hand sensorimotor cortex during/just before foot movement in foot versus hand movement paradigms is thought to reflect hand movement inhibition during current/prospective movement of another effector. Behavior of mu during music listening with movement suppressed has yet to be determined. We recorded 32-channel EEG ( n = 17) during silence without movement, overt movement (foot/hand), and music listening without movement. Using an independent component analysis-based source equivalent dipole clustering technique, we identified three mu-related clusters, localized to left primary motor and right and midline premotor cortices. Right foot tapping was accompanied by mu enhancement in the left lateral source cluster, replicating previous work. Music listening was accompanied by similar mu enhancement in the left, as well as midline, clusters. We are the first, to our knowledge, to report, and also to source-resolve, music-related mu modulation in the absence of overt movements. Covert music-related motor activity has been shown to play a role in beat perception (Ross JM, Iversen JR, Balasubramaniam R. Neurocase 22: 558-565, 2016). Our current results show enhancement in somatotopically organized mu, supporting overt motor inhibition during beat perception. NEW & NOTEWORTHY We are the first to report music-related mu enhancement in the absence of overt movements and the first to source-resolve mu activity during music listening. We suggest that music-related mu modulation reflects overt motor inhibition during passive music listening. This work is relevant for the development of theories relating to the involvement of covert motor system activity for predictive beat perception.

It has been hypothesized that to perform sensorimotor transformations efficiently, somatosensory information being fed back to a particular motor circuit is organized in accordance with the mechanical loading patterns of the skin that result from the motor activity generated by that circuit. Rearrangements of sensory information to different motor circuits could in this respect constitute a key component of sensorimotor learning. We here explored whether the organization of tactile input from the plantar forepaw of the rat to cortical and striatal circuits is affected by a period of extensive sensorimotor training in a skilled reaching and grasping task. Our data show that the representation of tactile stimuli in terms of both temporal and spatial response patterns changes as a consequence of the training and that spatial changes particularly involve the primary motor cortex. Based on the observed reorganization, we propose that reshaping of the spatiotemporal representation of the tactile afference to motor circuits is an integral component of the learning process that underlies skill acquisition in reaching and grasping. NEW & NOTEWORTHY Sensorimotor transformations are fundamental to the function of the nervous system and determine how patterns of sensory input are converted into appropriate movements. We here investigated the extent to which experience-dependent processes can reshape the organization of somatosensory input feeding into cortico-basal ganglia motor structures. Our data point to a particularly important role for the primary motor cortex in the functional adaptions associated with skilled motor learning.

In neurotypical individuals, arm choice in reaching movements depends on expected biomechanical effort, expected success, and a handedness bias. Following a stroke, does arm choice change to account for the decreased motor performance, or does it follow a preinjury habitual preference pattern? Participants with mild-to-moderate chronic stroke who were right-handed before stroke performed reaching movements in both spontaneous and forced-choice blocks, under no-time, medium-time, and fast-time constraint conditions designed to modulate reaching success. Mixed-effects logistic regression models of arm choice revealed that expected effort predicted choices. However, expected success only strongly predicted choice in left-hemiparetic individuals. In addition, reaction times decreased in left-hemiparetic individuals between the no-time and the fast-time constraint conditions but showed no changes in right-hemiparetic individuals. Finally, arm choice in the no-time constraint condition correlated with a clinical measure of spontaneous arm use for right-, but not for left-hemiparetic individuals. Our results are consistent with the view that right-hemiparetic individuals show a habitual pattern of arm choice for reaching movements relatively independent of failures. In contrast, left-hemiparetic individuals appear to choose their paretic left arm more optimally: that is, if a movement with the paretic arm is predicted to be not successful in the upcoming movement, the nonparetic right arm is chosen instead. NEW & NOTEWORTHY Although we are seldom aware of it, we constantly make decisions to use one arm or the other in daily activities. Here, we studied whether these decisions change following stroke. Our results show that effort, success, and side of lesion determine arm choice in a reaching task: whereas left-paretic individuals modified their arm choice in response to failures in reaching the target, right-paretic individuals showed a pattern of choice independent of failures.

The influence of proprioceptive feedback on muscle activity during isometric tasks is the subject of conflicting studies. We performed an isometric knee extension task experiment based on two common clinical tests for mobility and flexibility. The task was carried out at four preset angles of the knee, and we recorded from five muscles for two different hip positions. We applied muscle synergy analysis using nonnegative matrix factorization on surface electromyograph recordings to identify patterns in the data that changed with internal knee angle, suggesting a link between proprioception and muscle activity. We hypothesized that such patterns arise from the way proprioceptive and cortical signals are integrated in neural circuits of the spinal cord. Using the MIIND neural simulation platform, we developed a computational model based on current understanding of spinal circuits with an adjustable afferent input. The model produces the same synergy trends as observed in the data, driven by changes in the afferent input. To match the activation patterns from each knee angle and position of the experiment, the model predicts the need for three distinct inputs: two to control a nonlinear bias toward the extensors and against the flexors, and a further input to control additional inhibition of rectus femoris. The results show that proprioception may be involved in modulating muscle synergies encoded in cortical or spinal neural circuits. NEW & NOTEWORTHY The role of sensory feedback in motor control when limbs are held in a fixed position is disputed. We performed a novel experiment involving fixed position tasks based on two common clinical tests. We identified patterns of muscle activity during the tasks that changed with different leg positions and then inferred how sensory feedback might influence the observations. We developed a computational model that required three distinct inputs to reproduce the activity patterns observed experimentally. The model provides a neural explanation for how the activity patterns can be changed by sensory feedback.

Nervous systems face a torrent of sensory inputs, including proprioceptive feedback. Signal integration depends on spatially and temporally coinciding signals. It is unclear how relative time delays affect multimodal signal integration from spatially distant sense organs. We measured transmission times and latencies along all processing stages of sensorimotor pathways in the stick insect leg muscle control system, using intra- and extracellular recordings. Transmission times of signals from load-sensing tibial and trochanterofemoral campaniform sensilla (tiCS, tr/fCS) to the premotor network were longer than from the movement-sensing femoral chordotonal organ (fCO). We characterized connectivity patterns from tiCS, tr/fCS, and fCO afferents to identified premotor nonspiking interneurons (NSIs) and motor neurons (MNs) by distinguishing short- and long-latency responses to sensory stimuli. Functional NSI connectivity depended on sensory context. The timeline of multisensory integration in the NSI network showed an early phase of movement signal processing and a delayed phase of load signal integration. The temporal delay of load signals relative to movement feedback persisted into MN activity and muscle force development. We demonstrate differential delays in the processing of two distinct sensory modalities generated by the sensorimotor network and affecting motor output. The reported temporal differences in sensory processing and signal integration improve our understanding of sensory network computation and function in motor control. NEW & NOTEWORTHY Networks integrating multisensory input face the challenge of not only spatial but also temporal integration. In the local network controlling insect leg movements, proprioceptive signal delays differ between sensory modalities. Specifically, signal transmission times to and neuronal connectivity within the sensorimotor network lead to delayed information about leg loading relative to movement signals. Temporal delays persist up to the level of the motor output, demonstrating its relevance for motor control.

Action choices are influenced by future and recent past action states. For example, when performing two actions in succession, response times (RTs) to initiate the second action are reduced when the same hand is used. These findings suggest the existence of effector-specific processing for action planning. However, given that each hand is primarily controlled by the contralateral hemisphere, the RT benefit might actually reflect effector-independent, hemisphere-specific rather than effector-specific repetition effects. Here, participants performed two consecutive movements, each with a hand or a foot, in one of two directions. Direction was specified in an egocentric reference frame (inward, outward) or in an allocentric reference frame (left, right). Successive actions were initiated faster when the same limb (e.g., left hand-left hand), but not the other limb of the same body side (e.g., left foot-left hand), executed the second action. The same-limb advantage was evident even when the two movements involved different directions, whether specified egocentrically or allocentrically. Corroborating evidence from computational modeling lends support to the claim that repetition effects in action planning reflect persistent changes in baseline activity within neural populations that encode effector-specific action plans. NEW & NOTEWORTHY Repeated hand use facilitates the initiation of successive actions (repetition effect). This finding has been interpreted as evidence for effector-specific action plans. However, given that each hand is primarily controlled by the contralateral hemisphere, any differences might reflect effector-independent, hemisphere-specific rather than effector-specific processing. We dissociated these alternatives by asking participants to perform successive actions with hands and feet and provide novel evidence that repetition effects in limb use truly reflect effector-specific coding.

Spinal cord injury (SCI) commonly results in permanent loss of motor, sensory, and autonomic function. Recent clinical studies have shown that epidural spinal cord stimulation may provide a beneficial adjunct for restoring lower extremity and other neurological functions. Herein, we review the recent clinical advances of lumbosacral epidural stimulation for restoration of sensorimotor function in individuals with motor complete SCI and we discuss the putative neural pathways involved in this promising neurorehabilitative approach. We focus on three main sections: review recent clinical results for locomotor restoration in complete SCI; discuss the contemporary understanding of electrical neuromodulation and signal transduction pathways involved in spinal locomotor networks; and review current challenges of motor system modulation and future directions toward integrative neurorestoration. The current understanding is that initial depolarization occurs at the level of large diameter dorsal root proprioceptive afferents that when integrated with interneuronal and latent residual supraspinal translesional connections can recruit locomotor centers and augment downstream motor units. Spinal epidural stimulation can initiate excitability changes in spinal networks and supraspinal networks. Different stimulation parameters can facilitate standing or stepping, and it may also have potential for augmenting myriad other sensorimotor and autonomic functions. More comprehensive investigation of the mechanisms that mediate the transformation of dysfunctional spinal networks to higher functional states with a greater focus on integrated systems-based control system may reveal the key mechanisms underlying neurological augmentation and motor restoration after severe paralysis.

Although beta-band activity during motor planning is known to be modulated by uncertainty about where to act, less is known about its modulations to uncertainty about how to act. To investigate this issue, we recorded oscillatory brain activity with EEG while human participants ( n = 17) performed a hand choice reaching task. The reaching hand was either predetermined or of participants' choice, and the target was close to one of the two hands or at about equal distance from both. To measure neural activity in a motion artifact-free time window, the location of the upcoming target was cued 1,000-1,500 ms before the presentation of the target, whereby the cue was valid in 50% of trials. As evidence for motor planning during the cuing phase, behavioral observations showed that the cue affected later hand choice. Furthermore, reaction times were longer in the choice trials than in the predetermined trials, supporting the notion of a competitive process for hand selection. Modulations of beta-band power over central cortical regions, but not alpha-band or theta-band power, were in line with these observations. During the cuing period, reaches in predetermined trials were preceded by larger decreases in beta-band power than reaches in choice trials. Cue direction did not affect reaction times or beta-band power, which may be due to the cue being invalid in 50% of trials, retaining effector uncertainty during motor planning. Our findings suggest that effector uncertainty modulates beta-band power during motor planning. NEW & NOTEWORTHY Although reach-related beta-band power in central cortical areas is known to modulate with the number of potential targets, here we show, using a cuing paradigm, that the power in this frequency band, but not in the alpha or theta band, is also modulated by the uncertainty of which hand to use. This finding supports the notion that multiple possible effector-specific actions can be specified in parallel up to the level of motor preparation.

Adapting hand movements to changes in our body or the environment is essential for skilled motor behavior, as is the ability to flexibly combine experience gathered in separate contexts. However, it has been shown that when adapting hand movements to two different visuomotor perturbations in succession, interference effects can occur. Here, we investigate whether these interference effects compromise our ability to adapt to the superposition of the two perturbations. Participants tracked with a joystick, a visual target that followed a smooth but an unpredictable trajectory. Four separate groups of participants (total n = 83) completed one block of 50 trials under each of three mappings: one in which the cursor was rotated by 90° (ROTATION), one in which the cursor mimicked the behavior of a mass-spring system (SPRING), and one in which the SPRING and ROTATION mappings were superimposed (SPROT). The order of the blocks differed across groups. Although interference effects were found when switching between SPRING and ROTATION, participants who performed these blocks first performed better in SPROT than participants who had no prior experience with SPRING and ROTATION (i.e., composition). Moreover, participants who started with SPROT exhibited better performance under SPRING and ROTATION than participants who had no prior experience with each of these mappings (i.e., decomposition). Additional analyses confirmed that these effects resulted from components of learning that were specific to the rotational and spring perturbations. These results show that interference effects do not preclude the ability to compose/decompose various forms of visuomotor adaptation. NEW & NOTEWORTHY The ability to compose/decompose task representations is critical for both cognitive and behavioral flexibility. Here, we show that this ability extends to two forms of visuomotor adaptation in which humans have to perform visually guided hand movements. Despite the presence of interference effects when switching between visuomotor maps, we show that participants are able to flexibly compose or decompose knowledge acquired in previous sessions. These results further demonstrate the flexibility of sensorimotor adaptation in humans.

Many goal-directed actions that require rapid visuomotor planning and perceptual decision-making are affected in older adults, causing difficulties in execution of many functional activities of daily living. Visuomotor planning and perceptual identification are mediated by the dorsal and ventral visual streams, respectively, but it is unclear how age-induced changes in sensory processing in these streams contribute to declines in visuomotor decision-making performance. Previously, we showed that in young adults, task demands influenced movement strategies during visuomotor decision-making, reflecting differential integration of sensory information between the two streams. Here, we asked the question if older adults would exhibit deficits in interactions between the two streams during demanding motor tasks. Older adults ( n = 15) and young controls ( n = 26) performed reaching or interception movements toward virtual objects. In some blocks of trials, participants also had to select an appropriate movement goal based on the shape of the object. Our results showed that older adults corrected fewer initial decision errors during both reaching and interception movements. During the interception decision task, older adults made more decision- and execution-related errors than young adults, which were related to early initiation of their movements. Together, these results suggest that older adults have a reduced ability to integrate new perceptual information to guide online action, which may reflect impaired ventral-dorsal stream interactions. NEW & NOTEWORTHY Older adults show declines in vision, decision-making, and motor control, which can lead to functional limitations. We used a rapid visuomotor decision task to examine how these deficits may interact to affect task performance. Compared with healthy young adults, older adults made more errors in both decision-making and motor execution, especially when the task required intercepting moving targets. This suggests that age-related declines in integrating perceptual and motor information may contribute to functional deficits.

Motor skill learning involves improvement in feedforward control, the ability to execute a motor plan more reliably, and feedback control, the ability to adjust the motor plan on the fly. The dependence between these control components and the association between training conditions and their improvement have not been directly examined. This study characterizes the contribution of feedforward and feedback control components to motor skill learning using the arc-pointing task (APT), a drawing task that requires high motor acuity. In experiment 1 , the performance of three groups of subjects was tested before and after training with online visual feedback (OF group), with knowledge of performance feedback that was presented after movement completion (KP group), and with both online and KP feedback (KP + OF group). Although the improvement of the OF group was not different from the improvement of the KP + OF group, comparison of the KP and KP + OF groups revealed an advantage to the KP group in the fast test speed, suggesting that training without online feedback leads to a greater improvement in feedforward control. In experiment 2 , subject's improvement was examined using test probes for estimating feedback and feedforward control. Both KP + OF and KP groups showed improvement in feedforward and feedback conditions with a trend toward a greater improvement of the KP group. Our results suggest that online visual feedback suppresses improvement in feedforward control during motor skill learning. NEW & NOTEWORTHY Becoming a skillful player requires both executing reliable movements and being able to efficiently control them online. We study here how training with and without online visual feedback affects feedforward and feedback control improvement in a drawing task that requires high precision. We show that training with online feedback suppresses improvement in feedforward control and leads to inferior performance in fast movements.

Xenopus laevis has a lateral line mechanosensory system throughout its full life cycle, and a previous study on prefeeding stage tadpoles revealed that it may play a role in motor responses to both water suction and water jets. Here, we investigated the physiology of the anterior lateral line system in newly hatched tadpoles and the motor outputs induced by its activation in response to brief suction stimuli. High-speed videoing showed tadpoles tended to turn and swim away when strong suction was applied close to the head. The lateral line neuromasts were revealed by using DASPEI staining, and their inactivation with neomycin eliminated tadpole motor responses to suction. In immobilized preparations, suction or electrically stimulating the anterior lateral line nerve reliably initiated swimming but the motor nerve discharges implicating turning was observed only occasionally. The same stimulation applied during ongoing fictive swimming produced a halting response. The anterior lateral line nerve showed spontaneous afferent discharges at rest and increased activity during stimulation. Efferent activities were only recorded during tadpole fictive swimming and were largely synchronous with the ipsilateral motor nerve discharges. Finally, calcium imaging identified neurons with fluorescence increase time-locked with suction stimulation in the hindbrain and midbrain. A cluster of neurons at the entry point of the anterior lateral line nerve in the dorsolateral hindbrain had the shortest latency in their responses, supporting their potential sensory interneuron identity. Future studies need to reveal how the lateral line sensory information is processed by the central circuit to determine tadpole motor behavior. NEW & NOTEWORTHY We studied Xenopus tadpole motor responses to anterior lateral line stimulation using high-speed videos, electrophysiology and calcium imaging. Activating the lateral line reliably started swimming. At high stimulation intensities, turning was observed behaviorally but suitable motor nerve discharges were seen only occasionally in immobilized tadpoles. Suction applied during swimming produced a halting response. We analyzed afferent and efferent activities of the tadpole anterior lateral line nerve and located sensory interneurons using calcium imaging.

Well-documented manual dexterity impairments in older adults may critically depend on the processing of visual information. The purpose of this study was to determine age-related changes in eye and hand movements during commonly used pegboard tests and the association with manual dexterity impairments in older adults. The relationship between attentional deficits and manual dexterity was also assessed. Eye movements and hand kinematics of 20 young (20-38 yr) and 20 older (65-85 yr) adults were recorded during 9-Hole Pegboard, Grooved Pegboard, and a visuospatial dual test. Results were compared with standardized tests of attention (The Test of Everyday Attention and Trail Making Test) that assess visual selective attention, sustained attention, attentional switching, and divided attention. Hand movement variability was 34% greater in older versus young adults when placing the pegs into the pegboard and this was associated with decreased pegboard performance, providing further evidence that increased movement variability plays a role in dexterity impairments in older adults. Older adults made more corrective saccades and spent less time gazing at the pegboard than young adults, suggesting altered visual strategies in older compared with young adults. The relationship between pegboard completion time and Trail Making Test B demonstrates an association between attentional deficits and age-related pegboard impairments. Results contribute novel findings of age-associated changes in eye movements during a commonly used manual dexterity task and offer insight into potential mechanisms underlying hand motor impairments in older adults. NEW & NOTEWORTHY This eye tracking study contributes novel findings of age-associated changes in eye movements during the commonly used pegboard tests of manual dexterity, including a greater number of corrective saccades and lesser time gazing at the pegboard holes in older compared with young adults. An association between attentional deficits and dexterity impairments in older adults is also highlighted. Results shed light on potential mechanisms underlying well-documented motor deficits in older adults.

Human cerebral cortex can produce visuomotor responses that are modulated by contextual and task-specific constraints. However, the distributed cortical network for visuomotor transformations limits the minimal response time of that pathway. Notably, humans can generate express visuomotor responses in arm muscles that are inflexibly tuned to the target location and occur 80-120 ms from stimulus presentation [stimulus-locked responses (SLRs)]. This suggests a subcortical pathway for visuomotor transformations that might involve the superior colliculus and its downstream reticulo-spinal projections. Here we investigated whether cognitive expectations can modulate the SLR. In one experiment, we recorded surface electromyogram (EMG) from shoulder muscles as participants reached toward a visual target whose location was unpredictable in control conditions and partially predictable in cue conditions by interpreting a symbolic cue (75% validity). Valid symbolic cues led to earlier and larger SLRs than control conditions; invalid symbolic cues produced later and smaller SLRs than control conditions. This is consistent with a cortical top-down modulation of the putative subcortical SLR network. In a second experiment, we presented high-contrast targets in isolation (control) or ∼24 ms after low-contrast stimuli, which could appear at the same (valid cue) or opposite (invalid cue) location as the target and with equal probability (50% cue validity). We observed earlier SLRs than control with the valid low-contrast cues, whereas the invalid cues led to the opposite results. These findings may reflect bottom-up attentional mechanisms, potentially evolving subcortically via the superior colliculus. Overall, our results support both top-down and bottom-up modulations of the putative subcortical SLR network in humans. NEW & NOTEWORTHY Express visuomotor responses in humans appear to reflect subcortical sensorimotor transformation of visual inputs, potentially conveyed via the tecto-reticulo-spinal pathway. Here we show that the express responses are influenced by both symbolic and barely detectable spatial cues about stimulus location. The symbolic cue-induced effects suggest cortical top-down modulation of the putative subcortical visuomotor network. The effects of barely detectable cues may reflect exogenous facilitation mechanisms of the tecto-reticulo-spinal pathway.

Learning in visuomotor adaptation tasks is the result of both explicit and implicit processes. Explicit processes, operationalized as reaiming an intended movement to a new goal, account for a significant proportion of learning. However, implicit processes, operationalized as error-dependent learning that gives rise to aftereffects, appear to be highly constrained. The limitations of implicit learning are highlighted in the mirror-reversal task, where implicit corrections act in opposition to performance. This is surprising given the mirror-reversal task has been viewed as emblematic of implicit learning. One potential issue not being considered in these studies is that both explicit and implicit processes were allowed to operate concurrently, which may interact, potentially in opposition. Therefore, we sought to further characterize implicit learning in a mirror-reversal task with a clamp design to isolate implicit learning from explicit strategies. We confirmed that implicit adaptation is in the wrong direction for mirror reversal and operates as if the perturbation were a rotation and only showed a moderate attenuation after 3 days of training. This result raised the question of whether implicit adaptation blindly operates as though perturbations were a rotation. In a separate experiment, which directly compared a mirror reversal and a rotation, we found that implicit adaptation operates in a proper coordinate system for different perturbations: adaptation to a mirror reversal and rotational perturbation is more consistent with Cartesian and polar coordinate systems, respectively. It remains an open question why implicit process would be flexible to the coordinate system of a perturbation but continue to be directed inappropriately. NEW & NOTEWORTHY Recent studies have found that implicit learning may operate inappropriately in some motor tasks, requiring explicit strategies to improve performance. However, this inappropriate adaptation could be attributable to competitive interactions between explicit and implicit processes. After isolating implicit processes, we found that implicit adaptation remained in the wrong direction for a mirror reversal, acting as if it were a rotation. Interestingly, however, the implicit system is sensitive to a particular coordinate system, treating mirror reversal and rotation differently.

Postural muscle activity precedes voluntary movements of the upper limbs. The traditional view of this activity is that it anticipates perturbations to balance caused by the movement of a limb. However, findings from reach-based paradigms have shown that postural adjustments can initiate center of mass displacement for mobility rather than minimize its displacement for stability. Within this context, altering reaching distance beyond the base of support would place increasing constraints on equilibrium during stance. If the underlying composition of anticipatory postural activity is linked to stability, coordination between muscles (i.e., motor modules) may evolve differently as equilibrium constraints increase. We analyzed the composition of motor modules in functional trunk muscles as participants performed multidirectional reaching movements to targets within and beyond the arm's length. Bilateral trunk and reaching arm muscle activity were recorded. Despite different trunk requirements necessary for successful movement, and the changing biomechanical (i.e., postural) constraints that accompany alterations in reach distance, nonnegative matrix factorization identified functional motor modules derived from preparatory trunk muscle activity that shared common features. Relative similarity in modular weightings (i.e., composition) and spatial activation profiles that reflect movement goals across tasks necessitating differing levels of trunk involvement provides evidence that preparatory postural adjustments are linked to the same task priorities (i.e., movement generation rather than stability). NEW & NOTEWORTHY Reaching within and beyond arm's length places different task constraints upon the required trunk motion necessary for successful movement execution. The identification of constant modular features, including functional muscle weightings and spatial tuning, lend support to the notion that preparatory postural adjustments of the trunk are tied to the same task priorities driving mobility, regardless of the future postural constraints.

We investigated the role of task constraints on interpersonal interactions. Twenty-one pairs of coworkers performed a finger force production task on force sensors placed at two ends of a seesaw-like apparatus and matched a combined target force of 20 N for 23 s over 10 trials. There were two experimental conditions: 1 ) FIXED: the seesaw apparatus was mechanically held in place so that the only task constraint was to match the 20 N resultant force, and 2 ) MOVING: the lever in the apparatus was allowed to rotate freely around its fulcrum, acting like a seesaw, so an additional task constraint to (implicitly) balance the resultant moment was added. We hypothesized that the additional task constraint of moment stabilization imposed on the MOVING condition would deteriorate task performance compared with the FIXED condition; however, this was rejected, as the performance of the force matching task was similar between two conditions. We also hypothesized that the central nervous systems (CNSs) would employ distinct coworking strategies or interpersonal motor synergy (IPMS) between conditions to satisfy different task constraints, which was supported by our results. Negative covariance between coworker's forces in the FIXED condition suggested a force stabilization strategy, whereas positive covariance in the MOVING condition suggested a moment stabilization strategy, implying that independent CNSs adopt distinct IPMSs depending on task constraints. We speculate that in the absence of a central neural controller, shared visual and mechanical connections between coworkers may suffice to trigger modulations in the cerebellum of each CNS to satisfy competing task constraints. NEW & NOTEWORTHY To the best of our knowledge, this is the first study to investigate the coworking behavior or IPMS when an additional task constraint is imposed. Our proposed analytical framework quantifies IPMS and allows for investigating variability in offline (i.e., across multiple repetitions) and online (i.e., across time) control, which is novel in coworking research. Understanding variability while performing a task is essential, as repeating a task is not always possible, as in therapeutic contexts.

Different neurophysiological strategies are used to perform angle adjustments during motor tasks such as car driving and force-control tasks using a fixed-rigid pedal. However, the difference in motor unit behavior in response to an increasing exerted force between tasks is unknown. This study aimed to investigate the difference in motor unit responsiveness on increasing force between force and position tasks. Twelve healthy participants performed ramp and hold contractions during ankle plantarflexion at 20% and 30% of the maximal voluntary contraction using a rigid pedal (force task) and a free pedal with an inertial load (position task). High-density surface electromyograms were recorded of the medial gastrocnemius muscle and decomposed into individual motor unit firing patterns. Ninety and hundred and nine motor units could be tracked between different target torques in each task. The mean firing rate increased and firing rate variability decreased on 10% maximal voluntary contraction force gain during both force and position tasks. There were no significant differences in these responses between the two tasks. Our results suggest that the motor unit firing rate is similarly regulated between force and position tasks in the medial gastrocnemius muscle with an increase in the exerted force. NEW & NOTEWORTHY Different neurophysiological strategies are used to perform a force control task and angle adjustment task. Our results showed that motor unit firing rate is similarly regulated between the two tasks in the medial gastrocnemius muscle with an increase in the exerted force. Although it is reported that position tasks contribute to early fatigue, it does not seem to be a particular problem for the increase in force.

Reaching movements performed from a crouched body posture require a shift of body weight from both arms to one arm. This situation has remained unexamined despite the analogous load requirements during step initiation and the many studies of reaching from a seated or standing posture. To determine whether the body weight shift involves anticipatory or exclusively reactive control, we obtained force plate records, hand kinematics, and arm muscle activity from 11 healthy right-handed participants. They performed reaching movements with their left and right arm in two speed contexts, "comfortable" and "as fast as possible," and two postural contexts, a less stable knees-together posture and a more stable knees-apart posture. Weight-shifts involved anticipatory postural actions (APAs) by the reaching and stance arms that were opposing in the vertical axis and aligned in the side-to-side axis similar to APAs by the legs for step initiation. Weight-shift APAs were correlated in time and magnitude, present in both speed contexts, more vigorous with the knees placed together, and similar when reaching with the dominant and nondominant arm. The initial weight-shift was preceded by bursts of muscle activity in the shoulder and elbow extensors (posterior deltoid and triceps lateral) of the reach arm and shoulder flexor (pectoralis major) of the stance arm, which indicates their causal role; leg muscles may have indirectly contributed but were not recorded. The strong functional similarity of weight-shift APAs during crouched reaching to human stepping and cat reaching suggests that they are a core feature of posture-movement coordination. NEW & NOTEWORTHY This work demonstrates that reaching from a crouched posture is preceded by bimanual anticipatory postural adjustments (APAs) that shift the body weight to the stance limb. Weight-shift APAs are more robust in an unstable body posture (knees together) and involve the shoulder and elbow extensors of the reach arm and shoulder flexor of the stance arm. This pattern mirrors the forelimb coordination of cats reaching and humans initiating a step.

With effects of aging, voluntary neural drive to the muscle, measured as motor unit (MU) firing rate, is lower in older adults during sustained isometric contractions compared with young adults, but differences remain unknown during limb movements. Therefore, our purpose was to compare MU firing rates during both isometric and shortening contractions between two adult age groups. We analyzed intramuscular electromyography of single-MU recordings in the anconeus muscle of young ( n = 8, 19-33 yr) and very old ( n = 13, 78-93 yr) male adults during maximal voluntary contractions (MVCs). In sustained isometric and muscle-shortening contractions during limb movement, MU trains were linked with elbow joint kinematic parameters throughout the contraction time course. The older group was 33% weaker and 10% slower during movements than the young group ( P < 0.01). In isometric contractions, median firing rates were 42% lower ( P < 0.01) in the older group (18 Hz) compared with the young group (31 Hz), but during shortening contractions firing rates were higher for both age groups and not statistically different between groups. As a function of contraction time, firing rates at MU recruitment threshold were 39% lower in the older group, but the firing rate decrease was attenuated threefold throughout shortening contraction compared with the young group. At the single-MU level, age-related differences during isometric contractions (i.e., pre-movement initiation) do not remain constant throughout movement that comprises greater effects of muscle shortening. Results indicate that neural drive is task dependent and during movement in older adults it is decreased minimally. NEW & NOTEWORTHY Changes of neural drive to the muscle with adult aging, measured as motor unit firing rates during limb movements, are unknown. Throughout maximal voluntary efforts we found that, in comparison with young adults, firing rates were lower during isometric contraction in older adults but not different during elbow extension movements. Despite the older group being ∼33% weaker across contractions, their muscles can receive neural drive during movements that are similar to that of younger adults.

The ability to adjust movements to changes in the environment declines with aging. This age-related decline is caused by the decline of explicit adjustments. However, implicit adaptation remains intact and might even be increased with aging. Since proprioceptive information has been linked to implicit adaptation, it might well be that an age-related decline in proprioceptive acuity might be linked to the performance of older adults in implicit adaptation tasks. Indeed, age-related proprioceptive deficits could lead to altered sensory integration with an increased weighting of the visual sensory-prediction error. Another possibility is that reduced proprioceptive acuity results in an increased reliance on predicted sensory consequences of the movement. Both these explanations led to our preregistered hypothesis: we expected a relation between the decline of proprioception and the amount of implicit adaptation across ages. However, we failed to support this hypothesis. Our results question the existence of reliability-based integration of visual and proprioceptive signals during motor adaptation. NEW & NOTEWORTHY Is proprioceptive acuity linked to amount of implicit motor adaptation across ages? The latter is larger in old compared with younger people? In light of reliability-based sensory integration, we hypothesized that this larger implicit adaptation was linked to an age-related lower reliability of proprioception. Over 2 experiments and 130 participants, we failed to find any evidence for this. We discussed alternative explanations for the increase in implicit adaptation with age and the validity of our proprioceptive assessment.

Parkinsonian motor deficits are associated with elevated inhibitory output from the basal ganglia (BG). However, several features of Parkinson's disease (PD) have not been accounted for by this simple "classical rate model" framework, including the observation in patients with PD that movements guided by external stimuli are less impaired than otherwise identical movements generated based on internal goals. Is this difference due to divergent processing within the BG itself or due to the recruitment of extra-BG pathways by sensory processing? In addition, surprisingly little is known about precisely when, in the sequence from selecting to executing movements, BG output is altered by PD. Here, we address these questions by recording activity in the substantia nigra pars reticulata (SNr), a key BG output nucleus, in hemiparkinsonian mice performing a well-controlled behavioral task requiring stimulus-guided and internally specified directional movements. We found that hemiparkinsonian mice exhibited a bias ipsilateral to the side of dopaminergic cell loss that was stronger when movements were internally specified rather than stimulus guided, consistent with clinical observations in patients with Parkinson's disease. We further found that changes in parkinsonian SNr activity during movement preparation were consistent with the ipsilateral behavioral bias, as well as its greater magnitude for internally specified movements. Although these findings are inconsistent with some aspects of the classical rate model, they are accounted for by a related "directional rate model" positing that SNr output phasically overinhibits motor output in a direction-specific manner. These results suggest that parkinsonian changes in BG output underlying movement preparation contribute to the greater deficit in internally specified than stimulus-guided movements. NEW & NOTEWORTHY Movements of patients with Parkinson's disease are often less impaired when guided by external stimuli than when generated based on internal goals. Whether this effect is due to distinct processing in the basal ganglia (BG) or due to compensation from other motor pathways is an open question with therapeutic implications. We recorded BG output in behaving parkinsonian mice and found that BG activity during movement preparation was consistent with the differences between these forms of movement.

How does the brain coordinate concurrent adaptation of arm movements and standing posture? From previous studies, the postural control system can use information about previously adapted arm movement dynamics to plan appropriate postural control; however, it is unclear whether postural control can be adapted and controlled independently of arm control. The present study addresses that question. Subjects practiced planar reaching movements while standing and grasping the handle of a robotic arm, which generated a force field to create novel perturbations. Subjects were divided into two groups, for which perturbations were introduced in either an abrupt or a gradual manner. All subjects adapted to the perturbations while reaching with their dominant (right) arm and then switched to reaching with their nondominant (left) arm. Previous studies of seated reaching movements showed that abrupt perturbation introduction led to transfer of learning between arms, but gradual introduction did not. Interestingly, in this study neither group showed evidence of transferring adapted control of arm or posture between arms. These results suggest primarily that adapted postural control cannot be transferred independently of arm control in this task paradigm. In other words, whole body postural movement planning related to a concurrent arm task is dependent on information about arm dynamics. Finally, we found that subjects were able to adapt to the gradual perturbation while experiencing very small errors, suggesting that both error size and consistency play a role in driving motor adaptation. NEW & NOTEWORTHY This study examined adaptation of arm and postural control to novel dynamics while standing and reaching and subsequent transfer between reaching arms. Neither arm nor postural control was transferred between arms, suggesting that postural planning is highly dependent on the concurrent arm movement.

During active object manipulation, the finger-object interactions give rise to complex fingertip skin deformations. These deformations are in turn encoded by the local tactile afferents and provide rich and behaviorally relevant information to the central nervous system. Most of the work studying the mechanical response of the finger to dynamic loading has been performed under a passive setup, thereby precisely controlling the kinematics or the dynamics of the loading. However, to identify aspects of the deformations that are relevant to online control during object manipulation, it is desirable to measure the skin response in an active setup. To that end, we developed a device that allows us to monitor finger forces, skin deformations, and kinematics during fine manipulation. We describe the device in detail and test it to precisely describe how the fingertip skin in contact with the object deforms during a simple vertical oscillation task. We show that the level of grip force directly influences the fingerpad skin strains and that the strain rates are substantial during active manipulation (norm up to 100%/s). The developed setup will enable us to causally relate sensory information, i.e. skin deformation, to online control, i.e. grip force adjustment, in future studies. NEW & NOTEWORTHY We present a novel device, a manipulandum, that enables to image the contact between the finger and the contact surface during active manipulation of the device. The device is tested in a simple vertical oscillation task involving 18 participants. We demonstrate that substantial surface skin strains take place at the finger-object interface and argue that those deformations provide essential information for grasp stability during object manipulation.

Faster movements are typically more variable-a speed-accuracy trade-off known as Fitts' law. Are movements that are initiated faster also more variable? Neurophysiological work has associated larger neural variability during motor preparation with longer reaction time (RT) and larger movement variability, implying that movement variability decreases with increasing RT. Here, we recorded over 30,000 reaching movements in 11 human participants who moved to visually cued targets. Half of the visual cues were accompanied by a beep to evoke a wide RT range in each participant. Results show that initial reach variability decreases with increasing RT, for voluntarily produced RTs up to ∼300 ms, whereas other kinematic aspects and endpoint accuracy remained unaffected. We conclude that movement preparation time determines initial movement variability. We suggest that the chosen movement preparation time reflects a trade-off between movement initiation and precision. NEW & NOTEWORTHY Fitts' law describes the speed-accuracy trade-off in the execution of human movements. We examined whether there is also a trade-off between movement planning time and initial movement precision. We show that shorter reaction times result in higher initial movement variability. In other words, movement preparation time determines movement variability.

Illusionary mirror visual feedback alters interhemispheric communication and influences cross-limb interactions. Combining forceful unimanual contractions with the mirror illusion is a convenient way to provoke robust alterations within ipsilateral motor networks. It is unknown, however, if the mirror illusion affects cross-limb fatigability. We examine this concept by comparing the ipsilateral and contralateral handgrip force and electromyographic (EMG) responses following unimanual fatigue with and without illusionary mirror visual feedback. Participants underwent three experimental sessions (mirror, no-mirror, and control), performing a unimanual fatigue protocol with and without illusionary mirror visual feedback. Maximal handgrip force and EMG activity were measured before and after each session for both hands during maximal unimanual and bimanual contractions. The associated EMG activity from the inactive forearm during unimanual contraction was also examined. The novel findings demonstrate greater relative fatigability during bimanual versus unimanual contraction following unimanual fatigue (-31.8% vs. -23.4%, P < 0.01) and the mirror illusion attenuates this difference (-30.3% vs. -26.3%, P = 0.169). The results show no evidence for a cross-over effect of fatigue with (+0.62%, -2.72%) or without (+0.26%, -2.49%) the mirror illusion during unimanual or bimanual contraction. The mirror illusion resulted in significantly lower levels of associated EMG activity in the contralateral forearm. There were no sex differences for any of the measures of fatigability. These results demonstrate that the mirror illusion influences contraction-dependent fatigue during maximal handgrip contractions. Alterations in facilitatory and inhibitory transcallosal drive likely explain these findings. NEW & NOTEWORTHY Illusionary mirror visual feedback is a promising clinical tool for motor rehabilitation, yet many features of its influence on motor output are unknown. We show that maximal bimanual force output is compromised to a greater extent than unimanual force output following unimanual fatigue, yet illusionary mirror visual feedback attenuates this difference. The mirror illusion also reduces the unintended EMG activity of the inactive, contralateral forearm during unimanual contraction.

Safe and successful motor performance relies on the ability to adapt to physiological and environmental change and retain what is learned. An open question is what factors maximize this retention? One overlooked factor is the degree to which balance is challenged during learning. We propose that the greater need for control and/or perceived threat of falling or injury associated with balance-challenging tasks increases the value assigned to maintaining a learned visuomotor mapping (i.e., the new relationship between visual input and motor output). And we propose that a greater-valued mapping is a more retainable mapping, as it serves to benefit future motor performance. Thus, we tested the hypothesis that challenging balance enhances motor memory, reflected by greater recall and faster relearning (i.e., savings). Four groups of participants adapted to a novel visuomotor mapping induced by prism lenses while performing a reaching or walking task, with and without an additional balance challenge. We found that challenging balance did not disrupt visuomotor adaptation during reaching or walking. We then probed recall and savings by having participants repeat the adaptation protocol 1 wk later. For reaching, we found evidence of initial recall, though neither group demonstrated savings upon reexposure to the prisms. In contrast, both walking groups demonstrated significant initial recall and savings. In addition, we found that challenging balance significantly enhanced savings during walking. Taken together, our results demonstrate the robustness of motor memories formed during walking and highlight the potential influence of balance control on sensorimotor learning. NEW & NOTEWORTHY Most everyday tasks challenge our balance. Yet, this aspect of daily motor behavior is often overlooked in adaptation paradigms. Here, we show that challenging balance does not impair sensorimotor adaptation during precision reaching and walking tasks. Furthermore, we show that challenging balance enhances savings of a learned visuomotor mapping during walking. These results provide evidence for the potential performance benefits associated with learning during unconstrained, naturalistic behaviors.

Successful interaction with the environment requires the dissociation of self-induced from externally induced sensory stimulation. Temporal proximity of action and effect is hereby often used as an indicator of whether an observed event should be interpreted as a result of own actions or not. We tested how the delay between an action (press of a touch bar) and an effect (onset of simulated self-motion) influences the processing of visually simulated self-motion in the ventral intraparietal area (VIP) of macaque monkeys. We found that a delay between the action and the start of the self-motion stimulus led to a rise of activity above the baseline activity before motion onset in a subpopulation of 21% of the investigated neurons. In the responses to the stimulus, we found a significantly lower sustained activity when the press of a touch bar and the motion onset were contiguous compared to the condition when the motion onset was delayed. We speculate that this weak inhibitory effect might be part of a mechanism that sharpens the tuning of VIP neurons during self-induced motion and thus has the potential to increase the precision of heading information that is required to adjust the orientation of self-motion in everyday navigational tasks. NEW & NOTEWORTHY Neurons in macaque ventral intraparietal area (VIP) are responding to sensory stimulation related to self-motion, e.g. visual optic flow. Here, we found that self-motion induced activation depends on the sense of agency, i.e., it differed when optic flow was perceived as self- or externally induced. This demonstrates that area VIP is well suited for study of the interplay between active behavior and sensory processing during self-motion.

Many individuals who undergo limb amputation experience persistent phantom limb pain (PLP), but the underlying mechanisms of PLP are unknown. The traditional hypothesis was that PLP resulted from maladaptive plasticity in sensorimotor cortex that degrades the neural representation of the missing limb. However, a recent study of individuals with upper limb amputations has shown that PLP is correlated with aberrant electromyographic (EMG) activity in residual muscles, posited to reflect a retargeting of efferent projections from a preserved representation of a missing limb. Here, we assessed EMG activity in a residual thigh muscle (vastus lateralis, VL) in patients with transfemoral amputations during cyclical movements of a phantom foot. VL activity on the amputated side was compared to that recorded on patients' intact side while they moved both the phantom and intact feet synchronously. VL activity in the patient group was also compared to a sample of control participants with no amputation. We show that phantom foot movement is associated with greater VL activity in the amputated leg than that seen in the intact leg as well as that exhibited by controls. The magnitude of residual VL activity was also positively related to ratings of PLP. These results show that phantom limb movement is associated with aberrant activity in a residual muscle after lower-limb amputation and provide evidence of a positive relationship between this activity and phantom limb pain. NEW & NOTEWORTHY This study is the first to assess residual muscle activity during movement of a phantom limb in individuals with lower limb amputations. We find that phantom foot movement is associated with aberrant recruitment of a residual thigh muscle and that this aberrant activity is related to phantom limb pain.

Vision in depth is distorted. A similar distortion can be observed for pointing to visual targets in depth. It has been suggested that pointing errors in depth reflect the visual distortion. Alternatively, pointing in depth might be guided by a prior that biases movements toward the natural grasping distance at which object manipulation is usually performed. To dissociate whether pointing is guided by distorted vision only or whether it takes into account a natural grasping distance prior, we adapted pointing movements. Participants received visual feedback about the success of their pointing once the movement was finished. We distorted the feedback to signal either that pointing was not far enough or in separate sessions that pointing was too far. Participants adapted to this artificial error by either extending or shortening their pointing movements. The generalization of pointing adaptation revealed a bias in movement planning that is inconsistent with pointing being guided only by distorted vision but with the involvement of knowledge about the natural grasping distance. Adaptation was strongest for pointing movements to a middle position that corresponds to the natural grasping distance and it was weakest for movements leading away from it. It has been demonstrated that pointing adaptation in depth changes visual perception (Volcic R, Fantoni C, Caudek C, Assad JA, Domini F. J Neurosci 33: 17081-17088, 2013). We also wondered how effects of pointing adaptation on visual space would generalize in depth. We found that adaptation changed visual space, but that this change was independent of the adaptation direction. NEW & NOTEWORTHY Which information guides pointing in 3D space? Inaccuracies of vision in depth generate the need for the sensorimotor system to rely on other information sources to optimally plan movement trajectories. Here, we implemented pointing adaptation experiments that could dissociate if the generalization of adaptation follows visual distortions or if it is informed by a "natural grasping distance" prior. Adaptation was strongest for movements toward the natural grasping distance, suggesting the latter hypothesis to be true.

Successful object manipulation requires anticipatory high-level control of finger positions and forces to prevent object slip and tilt. Unlike young adults, who efficiently scale grip forces (GFs) according to surface conditions, old adults were reported to exert excessive grip forces. In this study, we theoretically show how grip force economy depends on the modulation of the centers of pressure on opposing grip surfaces (ΔCoP) according to object properties. In a grasp-to-lift study with young and elderly participants, we investigated how the instruction to lift the object with efficient GF influences the anticipation of torques, ΔCoP and GF control during complex variations of mass distributions and surface properties. Provision of the explicit instruction to strive for force efficiency prompted both age groups to optimize their ΔCoP modulation, although to a lesser degree in the elderly, and also led to a refinement of torque anticipation for a right-sided weight distribution in the young, but not the elderly participants. Consequently, marked drops in GF levels resulted. Furthermore, participants enhanced ΔCoP modulation and lowered GF safety ratios in challenging surface conditions. Higher GF in the elderly was due to decreased skin-surface friction but also worse ΔCoP modulation for lateralized mass distributions when trying to be force efficient. In contrast, safety margins were not elevated in the elderly, suggesting preserved GF control. Our findings demonstrate how task goals influence high-level motor control of object manipulation differentially in young and elderly participants and highlight the necessity to control for both instructions and friction when investigating GF control. NEW & NOTEWORTHY Previous studies have shown that forces are covaried as a function of centers of pressure (CoPs) to exert adequate torques. Here, we demonstrate that force-efficient object manipulation requires the modulation of CoPs and show that providing the instruction to be force efficient and challenging surface conditions elicits a GF safety ratio reduction as well as an optimization of anticipatory CoP modulation and torques in the young and, to a lesser degree, in the elderly.

The central nervous system (CNS) may produce coordinated motor outputs via the combination of motor modules representable as muscle synergies. Identification of muscle synergies has hitherto relied on applying factorization algorithms to multimuscle electromyographic data (EMGs) recorded during motor behaviors. Recent studies have attempted to validate the neural basis of the muscle synergies identified by independently retrieving the muscle synergies through CNS manipulations and analytic techniques such as spike-triggered averaging of EMGs. Experimental data have demonstrated the pivotal role of the spinal premotor interneurons in the synergies' organization and the presence of motor cortical loci whose stimulations offer access to the synergies, but whether the motor cortex is also involved in organizing the synergies has remained unsettled. We argue that one difficulty inherent in current approaches to probing the synergies' neural basis is that the EMG generative model based on linear combination of synergies and the decomposition algorithms used for synergy identification are not grounded on enough prior knowledge from neurophysiology. Progress may be facilitated by constraining or updating the model and algorithms with knowledge derived directly from CNS manipulations or recordings. An investigative framework based on evaluating the relevance of neurophysiologically constrained models of muscle synergies to natural motor behaviors will allow a more sophisticated understanding of motor modularity, which will help the community move forward from the current debate on the neural versus nonneural origin of muscle synergies.