Late integration of vision and proprioception during perturbed reaches.

The motor system corrects rapidly, but selectively, for perturbations to ongoing reaching movements, depending on the constraints of the task. To account for such sophistication, it has been postulated that corrections are based on an estimated limb state that integrates all sensory changes caused by the perturbation, taking into account their processing delays. Here, we asked if information from different sensory modalities is integrated immediately or processed separately in the early phase of a response. We perturbed the estimated state of the limb with both unimodal and bimodal visual and proprioceptive perturbations without changing the actual limb state. For visual perturbations, a cursor representing the hand was shifted to the left or the right relative to the true hand location. For proprioceptive perturbations, the biceps or triceps muscles were vibrated, which induced illusory limb-state changes to the right or the left. In the bimodal condition, the perturbations to vision and proprioception were either congruent or incongruent in their directions. Response latencies show that it takes ∼100 ms longer to respond to unimodal visual perturbations than to unimodal proprioceptive perturbations. Responses to bimodal perturbations show that it takes an additional ∼100 ms beyond the response to unimodal visual perturbations for intermodal consistency to impact the response. These results suggest that visual and proprioceptive signals are initially processed separately for state estimation and only combined at the level of the limb's motor output, instead of being immediately integrated into a single state estimate of the limb.NEW & NOTEWORTHY Both visual and proprioceptive signals provide information about arm state during reaching. By perturbing the perceived, but not the actual, position of the hand in both modalities using visual disturbances and muscle vibration, we examined multimodal integration and state estimation during reaching. Our results suggest that the early reach corrections are based on separate state estimates from the two sensory modalities and only later are based on a combined state estimate.

Integration of proprioceptive and visual feedback during online control of reaching.

Visual and proprioceptive feedback both contribute to perceptual decisions, but it remains unknown how these feedback signals are integrated together or consider factors such as delays and variance during online control. We investigated this question by having participants reach to a target with randomly applied mechanical and/or visual disturbances. We observed that the presence of visual feedback during a mechanical disturbance did not increase the size of the muscle response significantly but did decrease variance, consistent with a dynamic Bayesian integration model. In a control experiment, we verified that vision had a potent influence when mechanical and visual disturbances were both present but opposite in sign. These results highlight a complex process for multisensory integration, where visual feedback has a relatively modest influence when the limb is mechanically disturbed, but a substantial influence when visual feedback becomes misaligned with the limb.NEW & NOTEWORTHY Visual feedback is more accurate, but proprioceptive feedback is faster. How should you integrate these sources of feedback to guide limb movement? As predicted by dynamic Bayesian models, the size of the muscle response to a mechanical disturbance was essentially the same whether visual feedback was present or not. Only under artificial conditions, such as when shifting the position of a cursor representing hand position, can one observe a muscle response from visual feedback.

Motor plan differs for young and older adults during similar movements.

Older adults exhibit altered activation of the agonist and antagonist muscles during goal-directed movements compared with young adults. However, it remains unclear whether the differential activation of the antagonistic muscles in older adults results from an impaired motor plan or an altered ability of the muscle to contract. The purpose of this study, therefore, was to determine whether the motor plan differs for young and older adults. Ten young (26.1 ± 4.3 yr, 4 women) and 16 older adults (71.9 ± 6.9 yr, 9 women) participated in the study. Participants performed 100 trials of fast goal directed movements with ankle dorsiflexion while we recorded the electromyographic activity of the primary agonist (tibialis anterior; TA) and antagonist (soleus; SOL) muscles. From those 100 trials we selected 5 trials in each of 3 movement end-point categories (fast, accurate, and slow). We investigated age-associated differences in the motor plan by quantifying the individual activity and coordination of the agonist and antagonist muscles. During similar movement end points, older adults exhibited similar activation of the agonist (TA) and antagonist (SOL) muscles compared with young adults. In addition, the coordination of the agonist and antagonist muscles (TA and SOL) was different between the two age groups. Specifically, older adults exhibited lower TA-SOL overlap (F1,23 = 41.2, P < 0.001) and greater TA-SOL peak EMG delay (F1,25 = 35.5, P < 0.001). This finding suggests that although subjects in both age groups displayed similar movement end points, they exhibited a different motor plan, as demonstrated by altered coordination between the agonist and antagonist muscles.NEW & NOTEWORTHY We aimed to determine whether the altered activation of muscles in older adults compared with young adults during fast goal-directed movements is related to an altered motor plan. For matched movements, there were differences in the coordination of antagonistic muscles but no differences in the individual activation of muscles. We provide novel evidence that the differential activation of muscles in older adults is related to an altered motor plan.

Social gaze cueing elicits facilitatory and inhibitory effects on movement execution when the model might act on an object.

Social cues, such as eye gaze and pointing fingers, can increase the prioritisation of specific locations for cognitive processing. A previous study using a manual reaching task showed that, although both gaze and pointing cues altered target prioritisation (reaction times [RTs]), only pointing cues affected action execution (trajectory deviations). These differential effects of gaze and pointing cues on action execution could be because the gaze cue was conveyed through a disembodied head; hence, the model lacked the potential for a body part (i.e., hands) to interact with the target. In the present study, the image of a male gaze model, whose gaze direction coincided with two potential target locations, was centrally presented. The model either had his arms and hands extended underneath the potential target locations, indicating the potential to act on the targets (Experiment 1), or had his arms crossed in front of his chest, indicating the absence of potential to act (Experiment 2). Participants reached to a target that followed a nonpredictive gaze cue at one of three stimulus onset asynchronies. RTs and reach trajectories of the movements to cued and uncued targets were analysed. RTs showed a facilitation effect for both experiments, whereas trajectory analysis revealed facilitatory and inhibitory effects, but only in Experiment 1 when the model could potentially act on the targets. The results of this study suggested that when the gaze model had the potential to interact with the cued target location, the model's gaze affected not only target prioritisation but also movement execution.

Generalizing the optic flow equalization control law to an asymmetrical person-plus-object system.

Visually guided action in humans occurs in part through the use of control laws, which are dynamical equations in which optical information modulates an actor's interaction with their environment. For example, humans locomote through the center of a corridor by equalizing the speed of optic flow across their left and right fields of view. This optic flow equalization control law relies on a crucial assumption: that the shape of the body relative to the eyes is laterally symmetrical. Humans engaging in tool use are often producing person-plus-object systems that are not laterally symmetrical, such as when they hold a tool, bag, or briefcase in one hand, or when they drive a vehicle. This experiment tests a new generalized control law for centered steering that accounts for asymmetries produced by external tool use. Participants held an asymmetrical bar and centered themselves within a virtual moving hallway while the speed of the virtual walls were systematically changed. The results demonstrate that humans engaging with an asymmetrical tool can (1) perceive the asymmetry of a person-plus-object system, (2) use that information to modulate the use of optic flow equalization control laws for centered steering, and (3) functionally incorporate the asymmetrical tool into their perception-action system to successfully navigate their environment.

Motor cortex hemodynamic response to goal-oriented and non-goal-oriented tasks in healthy subjects.

Background: Motor disorders are one of the world's major scourges, and neuromotor rehabilitation is paramount for prevention and monitoring plans. In this scenario, exercises and motor tasks to be performed by patients are crucial to follow and assess treatments' progression and efficacy. Nowadays, in clinical environments, quantitative assessment of motor cortex activities during task execution is rare, due to the bulkiness of instrumentation and the need for immobility during measurements [e.g., functional magnetic resonance imaging (MRI)]. Functional near-infrared spectroscopy (fNIRS) can contribute to a better understanding of how neuromotor processes work by measuring motor cortex activity non-invasively in freely moving subjects. Aim: Exploit fNIRS to measure functional activation of the motor cortex area during arm-raising actions. Design: All subjects performed three different upper limbs motor tasks: arm raising (non-goal-oriented), arm raising and grasping (goal oriented), and assisted arm raising (passive task). Each task was repeated ten times. The block design for each task was divided into 5 seconds of baseline, 5 seconds of activity, and 15 seconds of recovery. Population: Sixteen healthy subjects (11 males and 5 females) with an average (+/- standard deviation) of 37.9 (+/- 13.0) years old. Methods: Cerebral hemodynamic responses have been recorded in two locations, motor cortex (activation area) and prefrontal cortex (control location) exploiting commercial time-domain fNIRS devices. Haemodynamic signals were analyzed, separating the brain cortex hemodynamic response from extracerebral hemodynamic variations. Results: The hemodynamic response was recorded in the cortical motor area for goal-oriented and not-goaloriented tasks, while no response was noticed in the control location (prefrontal cortex position). Conclusions: This study provides a basis for canonical upper limb motor cortex activations that can be potentially compared to pathological cerebral responses in patients. It also highlights the potential use of TD-fNIRS to study goal-oriented versus non-goaloriented motor tasks. Impact: the findings of this study may have implications for clinical rehabilitation by providing a better understanding of the neural mechanisms underlying goal-oriented versus non-goal-oriented motor tasks. This may lead to more effective rehabilitation strategies for individuals with motor disorders and a more effective diagnosis of motor dysfunction supported by objective and quantitative neurophysiological readings.

Revealing the effects of temporal orienting of attention on response conflict using continuous movements.

Orienting attention in time enables us to prepare for forthcoming perception and action (e.g., estimating the duration of a yellow traffic light when driving). While temporal orienting can facilitate performance on simple tasks, its influence on complex tasks involving response conflict is unclear. Here, we adapted the flanker paradigm to a choice-reaching task where participants used a computer mouse to reach to the left or right side of the screen, as indicated by the central arrow presented with either the congruent or incongruent flankers. We assessed the effects of temporal orienting by manipulating goal-driven temporal expectation (using probabilistic variations in target timing) and stimulus-driven temporal priming (using sequential repetitions versus switches in target timing). We tested how temporal orienting influenced the dynamics of response conflict resolution. Recent choice-reaching studies have indicated that under response conflict, delayed movement initiation captures the response threshold adjustment process, whereas increased curvature toward the incorrect response captures the degree of coactivation of the response alternatives during the controlled response selection process. Both temporal expectation and priming reduced the initiation latency regardless of response conflict, suggesting that both lowered response thresholds independently of response conflict. Notably, temporal expectation, but not temporal priming, increased the curvature toward the incorrect response on incongruent trials. These results suggest that temporal orienting generally increases motor preparedness, but goal-driven temporal orienting particularly interferes with response conflict resolution, likely through its influence on response thresholds. Overall, our study highlights the interplay between temporal orienting and cognitive control in goal-directed action.

The effects of testing environment, experimental design, and ankle loading on calibration to perturbed optic flow during locomotion.

Calibration is the process by which the execution of actions becomes scaled to the (changing) relationship between environmental features and the actor's action capabilities. Though much research has investigated how individuals calibrate to perturbed optic flow, it remains unclear how different experimental factors contribute to the magnitude of calibration transfer. In the present study, we assessed how testing environment (Experiment 1), an adapted pretest-calibration-posttest design (Experiment 2), and bilateral ankle loading (Experiment 3) affected the magnitude of calibration to perturbed optic flow. We found that calibration transferred analogously to real-world and virtual environments. Although the magnitude of calibration transfer found here was greater than that reported by previous researchers, it was evident that calibration occurred rapidly and quickly plateaued, further supporting the claim that calibration is often incomplete despite continued calibration trials. We also saw an asymmetry in calibration magnitude, which may be due to a lack of appropriate perceptual-motor scaling prior to calibration. The implications of these findings for the assessment of distance perception and calibration in real-world and virtual environments are discussed.

Visual selection and response selection without effector selection in tasks with circular arrays.

Charles Eriksen and colleagues conducted influential visual-search experiments with circular arrays for which the responses were either vocal naming or unimanual left-right switch movements. These methods have the advantages of the stimuli being equidistant from a centered fixation point and allowing study of visual selection and response selection when effector selection is not required, as in the more typical case in which responses are key presses of distinct fingers. Other researchers have used similar spatial arrangements, but with aimed movements of the limb or of a mouse-controlled cursor to study effects of stimulus identification, visual search, spatial stimulus-response compatibility, response-effect compatibility, and practice/transfer in isolation and jointly. We systematically review studies in these areas that include visual selection and response selection and execution, and examine implications of their results for the role of effector selection. Also, we illustrate that as one moves from simpler to more complex tasks, the results are consistent with a basic information-processing framework in which stimulus identification and selection of a target response location are distinct from selecting, planning, and moving an effector to the targeted location.

Reaching movements are attracted by stimuli that signal reward.

When presented with a set of possible reach targets, the movement trajectory can reveal aspects of the underlying competition for action selection. Current goals and physical salience can affect the trajectory of reaching movements to be attracted towards a distractor. Some studies demonstrated that stimuli associated with reward can also cause an attraction when reaching towards the reward stimulus was previously rewarded and the reward stimulus was physically salient. Here we demonstrate that a non-salient stimulus that signals the availability of reward attracts reaching movements even when moving towards it was never necessary nor rewarded. Moreover, the attraction by reward is particularly evident with short-latency movements. We conclude that neither physical salience nor reinforcing the movement towards a stimulus is necessary for reward to gain priority in the selection for action.

Symbolic, non-directional predictive cues affect action execution.

Predictive cues may help us to plan an action in anticipation of what will come next. Some cues such as arrows contain directional information to orient the actor. Other cues, however, may contain no spatial information that directly orients the actor to the upcoming action. Non-directional predictive cues have been shown to increase performance in visual search tasks but have not been explored in the planning and execution of actions. The first aim of this study is to determine whether participants can implicitly learn to associate symbolic cues with an upcoming action target location. The second aim is whether this association leads to transient or sustained activation of the action associated with the predicted target location. High and low predictive cues preceded target appearance at long cue-target-onset asynchronies (1,100-2,000 ms). The trajectories of participants' reaching movements were analyzed depending on whether they aimed at the predicted or the non-predicted side within each of the cue type conditions. For the highly predictive cue, participants' trajectories veered further toward the opposite target location when participants aimed for the predicted target location compared to when they aimed for the non-predicted target location. These results indicate that participants can associate an upcoming action with non-directional predictive cues but the predicted response was inhibited in the long intervening time between the cue and target. This finding is similar to the response to peripheral-onset cues in inhibition of return type paradigms.

Computer mouse tracking reveals motor signatures in a cognitive task of spatial language grounding.

In a novel computer mouse tracking paradigm, participants read a spatial phrase such as "The blue item to the left of the red one" and then see a scene composed of 12 visual items. The task is to move the mouse cursor to the target item (here, blue), which requires perceptually grounding the spatial phrase. This entails visually identifying the reference item (here, red) and other relevant items through attentional selection. Response trajectories are attracted toward distractors that share the target color but match the spatial relation less well. Trajectories are also attracted toward items that share the reference color. A competing pair of items that match the specified colors but are in the inverse spatial relation increases attraction over-additively compared to individual items. Trajectories are also influenced by the spatial term itself. While the distractor effect resembles deviation toward potential targets in previous studies, the reference effect suggests that the relevance of the reference item for the relational task, not its role as a potential target, was critical. This account is supported by the strengthened effect of a competing pair. We conclude, therefore, that the attraction effects in the mouse trajectories reflect the neural processes that operate on sensorimotor representations to solve the relational task. The paradigm thus provides an experimental window through motor behavior into higher cognitive function and the evolution of activation in modal substrates, a longstanding topic in the area of embodied cognition.

Probing the time course of facilitation and inhibition in gaze cueing of attention in an upper-limb reaching task.

Previous work has revealed that social cues, such as gaze and pointed fingers, can lead to a shift in the focus of another person's attention. Research investigating the mechanisms of these shifts of attention has typically employed detection or localization button-pressing tasks. Because in-depth analyses of the spatiotemporal characteristics of aiming movements can provide additional insights into the dynamics of the processing of stimuli, in the present study we used a reaching paradigm to further explore the processing of social cues. In Experiments 1 and 2, participants aimed to a left or right location after a nonpredictive eye gaze cue toward one of these target locations. Seven stimulus onset asynchronies (SOAs), from 100 to 2,400 ms, were used. Both the temporal (reaction time, RT) and spatial (initial movement angle, IMA) characteristics of the movements were analyzed. RTs were shorter for cued (gazed-at) than for uncued targets across most SOAs. There were, however, no statistical differences in IMAs between movements to cued and uncued targets, suggesting that action planning was not affected by the gaze cue. In Experiment 3, the social cue was a finger pointing to one of the two target locations. Finger-pointing cues generated significant cueing effects in both RTs and IMAs. Overall, these results indicate that eye gaze and finger-pointing social cues are processed differently. Perception-action coupling (i.e., a tight link between the response and the social cue that is presented) might play roles in both the generation of action and the deviation of trajectories toward cued and uncued targets.

Endpoint accuracy of goal-directed ankle movements correlates to over-ground walking in stroke.

OBJECTIVES: Goal-directed movements are essential for voluntary motor control. The inability to execute precise goal-directed movements after stroke can impair the ability to perform voluntary functions, learn new skills, and hinder rehabilitation. However, little is known about how the accuracy of single-joint, goal-directed ankle movements relates to multi-joint, lower limb function in stroke. Here, we determined the impact of stroke on the accuracy of goal-directed ankle movements and its relation to over-ground walking. METHODS: Stroke (N = 28) and control (N = 28) participants performed (1) goal-directed ankle dorsiflexion movements to accurately match 9 degrees in 180 ms and (2) over-ground walking. During goal-directed ankle movements, we measured the endpoint error, position error, time error and the activation of the agonist and antagonist muscles. During over-ground walking, we measured the walking speed, paretic stride length, and cadence. RESULTS: The stroke group demonstrated increased endpoint error than the controls. Increased endpoint error was associated with increased co-activation between agonist-antagonist muscles. Endpoint error was a significant predictor of walking speed and paretic stride length in stroke. CONCLUSIONS: Impaired accuracy of goal-directed, ankle movements is correlated to over-ground walking in stroke. SIGNIFICANCE: Quantifying accuracy of goal-directed ankle movements may provide insights into walking function post-stroke.

Cholinergic Sensorimotor Integration Regulates Olfactory Steering.

Sensorimotor integration regulates goal-directed movements. We study the signaling mechanisms underlying sensorimotor integration in C. elegans during olfactory steering, when the sinusoidal movements of the worm generate an in-phase oscillation in the concentration of the sampled odorant. We show that cholinergic neurotransmission encodes the oscillatory sensory response and the motor state of head undulations by acting through an acetylcholine-gated channel and a muscarinic acetylcholine receptor, respectively. These signals converge on two axonal domains of an interneuron RIA, where the sensory-evoked signal suppresses the motor-encoding signal to transform the spatial information of the odorant into the asymmetry between the axonal activities. The asymmetric synaptic outputs of the RIA axonal domains generate a directional bias in the locomotory trajectory. Experience alters the sensorimotor integration to generate specific behavioral changes. Our study reveals how cholinergic neurotransmission, which can represent sensory and motor information in the mammalian brain, regulates sensorimotor integration during goal-directed locomotions.

Visual monitoring of goal-directed aiming movements.

Goal-directed movements are subject to intrinsic planning and execution variability, which requires that the central nervous system closely monitor our movements to ensure endpoint accuracy. In the present study, we sought to determine how closely the visual system monitored goal-directed aiming movements. We used a cursor-jump paradigm in which a cursor was unexpectedly translated soon after movement initiation. Some of the trials included a second cursor jump, and the cursor remained visible for different durations. The results indicate that seeing the cursor for only 16 ms after the second cursor jump was sufficient to influence the movement endpoint, which suggests that the visual system continuously monitored goal-directed movements. The results also suggest that the perceived position/trajectory of the effector was likely to have been averaged over a period of approximately 70 ms.

Reduced asymmetry in motor skill learning in left-handed compared to right-handed individuals.

Hemispheric specialization for motor control influences how individuals perform and adapt to goal-directed movements. In contrast to adaptation, motor skill learning involves a process wherein one learns to synthesize novel movement capabilities in absence of perturbation such that they are performed with greater accuracy, consistency and efficiency. Here, we investigated manual asymmetry in acquisition and retention of a complex motor skill that requires speed and accuracy for optimal performance in right-handed and left-handed individuals. We further determined if degree of handedness influences motor skill learning. Ten right-handed (RH) and 10 left-handed (LH) adults practiced two distinct motor skills with their dominant or nondominant arms during separate sessions two-four weeks apart. Learning was quantified by changes in the speed-accuracy tradeoff function measured at baseline and one-day retention. Manual asymmetry was evident in the RH group but not the LH group. RH group demonstrated significantly greater skill improvement for their dominant-right hand than their nondominant-left hand. In contrast, for the LH group, both dominant and nondominant hands demonstrated comparable learning. Less strongly-LH individuals (lower EHI scores) exhibited more learning of their dominant hand. These results suggest that while hemispheric specialization influences motor skill learning, these effects may be influenced by handedness.

Action plans can interact to hinder or facilitate reach performance.

Executing a reach action can be delayed while retaining another action in working memory (WM) if the two action plans partly overlap rather than do not overlap. This delay (partial repetition cost) occurs when reach responses are under cognitive control. In this study, we investigated whether facilitation (a partial repetition benefit) occurs when reach responses are automatic. We also examined whether the hemisphere controlling the limb or selection of the preferred limb (based on a free-reach task) influences reach performance when the actions partly overlap. Left- and right-handers reached to different stimulus locations to the left and right of body midline with their ipsilateral hand while maintaining an action plan in WM that required the same or the different hand. The results showed a partial repetition benefit for spatially compatible reaches to left and right stimulus locations far from the body midline, but not for those near the body midline. Also, no partial repetition cost was found at any of the stimulus-reach locations. This indicates that automatic reach responses that partly overlap with an action plan maintained in WM are not delayed, but instead can be facilitated (partial repetition benefit). The roles of hemisphere and reach-hand preference in action control and the importance of the degree of feature overlap in obtaining a partial repetition benefit (and cost) are discussed.