The effects of auditory consequences on visuomotor adaptation and motor memory.

Sensorimotor adaptation is a form of motor learning that is essential for maintaining motor performance across the lifespan and is integral to recovery of function after neurological injury. Recent research indicates that experiencing a balance-threatening physical consequence when making a movement error during adaptation can enhance subsequent motor memory. This is perhaps not surprising, as learning to avoid injury is critical for our survival and well-being. Reward and punishment can also differentially modify aspects of motor learning. However, it remains unclear whether other forms of non-physical consequences can impact motor learning. Here we tested the hypothesis that a loud acoustic stimulus linked to a movement error during adaptation could lead to greater generalization and consolidation. Two groups of participants (n = 12 each) adapted to a new, prism-induced visuomotor mapping while performing a precision walking task. One group experienced an unexpected loud acoustic stimulus (85 dB tone) when making foot-placement errors during adaptation. This auditory consequence group adapted faster and showed greater generalization with an interlimb transfer task, but not greater generalization to an obstacle avoidance task. Both groups showed faster relearning (i.e., savings) during the second testing session one week later despite the presence of an interference block of trials following initial adaptation, indicating successful consolidation. However, we did not find significant differences between groups with relearning during session 2. Overall, our results suggest that auditory consequences may serve as a useful method to improve motor learning, though further research is required.

Interactions during falls with environmental objects: evidence from real-life falls in long-term care captured on video.

BACKGROUND: Falls are the leading cause of injuries in older adults. Environmental objects (such as furniture, walls, and handrails) may act as hazards or facilitators to balance maintenance and safe landing. There is lack of objective evidence on how older adults interact with objects during falls. We addressed this gap by characterizing body part contacts with objects other than the floor during real-life falls in long-term care. METHODS: We analyzed videos of 1759 falls experienced by 584 residents to characterize the prevalence of contacts with objects before, during, and after fall initiation. Using generalized estimating equations, we compared the prevalence of falls with versus without contact to objects after fall initiation. Using linear mixed models, we tested for differences across body parts in the probability of contacting objects after fall initiation. RESULTS: In nearly one-third of falls, interactions with objects (e.g., trips over objects, loss of support with objects) or with other people (e.g., being pushed by another person) had a primary role in causing imbalance and initiating the fall. After fall initiation, participants contacted objects in 60% of falls, with intentional hand contacts to objects via reach-to-grasp or bracing being the most common type of interaction (Probability ± SE = 0.32 ± 0.01), followed by unintentional impacts to the torso (0.21 ± 0.01) and head (0.16 ± 0.01). Intentional hand contact to an object was more common during forward than backward falls (p < 0.001), while head and torso contacts to objects were more common during backward and sideways falls than forward falls (multiple p values ≤ 0.003). The hand most often contacted chairs, wheelchairs or couches, followed by tables or counters, walls, other people, walkers, and handrails. The head, torso, and shoulder most often contacted a wall. CONCLUSIONS: Most falls in long-term care involved contacts with objects other than the ground, indicating that complex environments often accompany falls in long-term care. Higher probabilities of intentional hand contacts in forward falls, versus unintentional head and torso impacts in backward and sideways falls may reflect the influence of being able to visualize and adjust one's falling patterns to nearby objects.

Fork in the road: How self-efficacy related to walking across terrain influences gaze behavior and path choice.

Decisions about where to move occur throughout the day and are essential to life. Different movements may present different challenges and affect the likelihood of achieving a goal. Certain choices may have unintended consequences, some of which may cause harm and bias the decision. Movement decisions rely on a person gathering necessary visual information via shifts in gaze. Here we sought to understand what influences this information-seeking gaze behavior. Participants chose between walking across one of two paths that consisted of terrain images found in either hiking or urban environments. We manipulated the number and type of terrain of each path, which altered the amount of available visual information. We recorded gaze behavior during the approach to the paths and had participants rate the confidence in their ability to walk across each terrain type (i.e., self-efficacy) as though it was real. Participants did not direct gaze more to the path with greater visual information, regardless of how we quantified information. Rather, we show that a person's perception of their motor abilities predicts how they visually explore the environment with their eyes as well as their choice of action. The greater the self-efficacy in walking across one path, the more they directed gaze to it and the more likely they chose to walk across it.

Savings in sensorimotor learning during balance-challenged walking but not reaching.

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.

Cortical Effects of Noisy Galvanic Vestibular Stimulation Using Functional Near-Infrared Spectroscopy.

Noisy galvanic vestibular stimulation (nGVS) can improve different motor, sensory, and cognitive behaviors. However, it is unclear how this stimulation affects brain activity to facilitate these improvements. Functional near-infrared spectroscopy (fNIRS) is inexpensive, portable, and less prone to motion artifacts than other neuroimaging technology. Thus, fNIRS has the potential to provide insight into how nGVS affects cortical activity during a variety of natural behaviors. Here we sought to: (1) determine if fNIRS can detect cortical changes in oxygenated (HbO) and deoxygenated (HbR) hemoglobin with application of subthreshold nGVS, and (2) determine how subthreshold nGVS affects this fNIRS-derived hemodynamic response. A total of twelve healthy participants received nGVS and sham stimulation during a seated, resting-state paradigm. To determine whether nGVS altered activity in select cortical regions of interest (BA40, BA39), we compared differences between nGVS and sham HbO and HbR concentrations. We found a greater HbR response during nGVS compared to sham stimulation in left BA40, a region previously associated with vestibular processing, and with all left hemisphere channels combined (p < 0.05). We did not detect differences in HbO responses for any region during nGVS (p > 0.05). Our results suggest that fNIRS may be suitable for understanding the cortical effects of nGVS.

Challenging balance during sensorimotor adaptation increases generalization.

From reaching to walking, real-life experience suggests that people can generalize between motor behaviors. One possible explanation for this generalization is that real-life behaviors often challenge our balance. We propose that the exacerbated body motions associated with balance-challenged whole body movements increase the value to the nervous system of using a comprehensive internal model to control the task. Because it is less customized to a specific task, a more comprehensive model is also a more generalizable model. Here we tested the hypothesis that challenging balance during adaptation would increase generalization of a newly learned internal model. We encouraged participants to learn a new internal model using prism lenses that created a new visuomotor mapping. Four groups of participants adapted to prisms while performing either a standing-based reaching or precision walking task, with or without a manipulation that challenged balance. To assess generalization after the adaptation phase, participants performed a single trial of each of the other groups' tasks without prisms. We found that both the reaching and walking balance-challenged groups showed significantly greater generalization to the equivalent, nonadapted task than the balance-unchallenged groups. Additionally, we found some evidence that all groups generalized across tasks, for example, from walking to reaching and vice versa, regardless of balance manipulation. Overall, our results demonstrate that challenging balance increases the degree to which a newly learned internal model generalizes to untrained movements.NEW & NOTEWORTHY Real-life experience indicates that people can generalize between motor behaviors. Here we show that challenging balance during the learning of a new internal model increases the degree of generalization to untrained movements for both reaching and walking tasks. These results suggest that the effects of challenging balance are not specific to the task but instead apply to motor learning more broadly.

Motor cost affects the decision of when to shift gaze for guiding movement.

Frequent gait modifications are often required to navigate our world. These can involve long or wide steps or changes in direction. People generally prefer to minimize the motor cost (or effort) of a movement, although with changes in gait this is not always possible. The decision of when and where to shift gaze is critical for controlling motor actions, since vision informs the brain about the available choices for movement-in this case, where to step. Here we asked how motor cost influences the allocation of gaze. To address this, we had participants walk and step to the center of sequential targets on the ground. We manipulated the motor cost associated with controlling foot placement by varying the location of one target in the lateral direction on a trial-to-trial basis within environments with different numbers of targets. Costlier steps caused a switch from a gaze strategy of planning future steps to one favoring visual feedback of the current foot placement when participants had to negotiate another target immediately after. Specifically, costlier steps delayed gaze shifts away from the manipulated target. We show that this relates to the cost of moving the leg and redirecting the body's center of mass from target to target. Overall, our results suggest that temporal gaze decisions are affected by motor costs associated with step-to-step demands of the environment. Moreover, they provide insight into what affects the coordination between the eyes and feet for the control of stable and accurate foot placement while walking. NEW & NOTEWORTHY Changes in gait allow us to navigate our world. For instance, one may step long or wide to avoid a spilled drink. The brain can direct gaze to gather relevant information for making these types of motor decisions; however, the factors affecting gaze allocation in natural behaviors are poorly understood. We show how the motor cost associated with a step influences the decision of when to redirect gaze to ensure accurate foot placement while walking.

Long-term retention and reconsolidation of a visuomotor memory.

Visuomotor adaptation is a form of motor learning that enables accurate limb movements in the presence of altered environmental or internal conditions. It requires updating the mapping between visual input and motor output, and can occur when learning a new device/tool or during rehabilitation after neurological injury. In either case, it is desirable to stabilize, or consolidate, this visuomotor memory for long-term usage. However, reactivation of a consolidated memory, whether it is motor-based or not, is thought to render it temporarily fragile again, and thus susceptible to interference or modification. Here, we determined if visuomotor memories demonstrate long-term retention but are fragile once reactivated. We used prism lenses to create a novel visuomotor mapping, which participants learned while having to walk and step to the center of targets. We re-tested this memory after one week and one year. We found that the mapping is retained for at least one year, regardless of whether participants were exposed to an interfering (i.e., opposing) mapping in the first session. We also found that presenting an opposing mapping in a block of trials following reactivation of the memory one year later did not disrupt subsequent performance when we re-tested the original memory. Our results suggest that these visuomotor memories are stored for extended periods of time and have limited fragility. Taken together, our results highlight the robustness of visuomotor memories associated with walking.

Consolidation of visuomotor adaptation memory with consistent and noisy environments.

Our understanding of how we learn and retain motor behaviors is still limited. For instance, there is conflicting evidence as to whether the memory of a learned visuomotor perturbation consolidates; i.e., the motor memory becomes resistant to interference from learning a competing perturbation over time. Here, we sought to determine the factors that influence consolidation during visually guided walking. Subjects learned a novel mapping relationship, created by prism lenses, between the perceived location of two targets and the motor commands necessary to direct the feet to their positions. Subjects relearned this mapping 1 wk later. Different groups experienced protocols with or without a competing mapping (and with and without washout trials), presented either on the same day as initial learning or before relearning on day 2 We tested identical protocols under constant and noisy mapping structures. In the latter, we varied, on a trial-by-trial basis, the strength of prism lenses around a non-zero mean. We found that a novel visuomotor mapping is retained at least 1 wk after initial learning. We also found reduced foot-placement error with relearning in constant and noisy mapping groups, despite learning a competing mapping beforehand, and with the exception of one protocol, with and without washout trials. Exposure to noisy mappings led to similar performance on relearning compared with the equivalent constant mapping groups for most protocols. Overall, our results support the idea of motor memory consolidation during visually guided walking and suggest that constant and noisy practices are effective for motor learning. NEW & NOTEWORTHY: The adaptation of movement is essential for many daily activities. To interact with targets, this often requires learning the mapping to produce appropriate motor commands based on visual input. Here, we show that a novel visuomotor mapping is retained 1 wk after initial learning in a visually guided walking task. Furthermore, we find that this motor memory consolidates (i.e., becomes more resistant to interference from learning a competing mapping) when learning in constant and noisy mapping environments.

Foot placement relies on state estimation during visually guided walking.

As we walk, we must accurately place our feet to stabilize our motion and to navigate our environment. We must also achieve this accuracy despite imperfect sensory feedback and unexpected disturbances. In this study we tested whether the nervous system uses state estimation to beneficially combine sensory feedback with forward model predictions to compensate for these challenges. Specifically, subjects wore prism lenses during a visually guided walking task, and we used trial-by-trial variation in prism lenses to add uncertainty to visual feedback and induce a reweighting of this input. To expose altered weighting, we added a consistent prism shift that required subjects to adapt their estimate of the visuomotor mapping relationship between a perceived target location and the motor command necessary to step to that position. With added prism noise, subjects responded to the consistent prism shift with smaller initial foot placement error but took longer to adapt, compatible with our mathematical model of the walking task that leverages state estimation to compensate for noise. Much like when we perform voluntary and discrete movements with our arms, it appears our nervous systems uses state estimation during walking to accurately reach our foot to the ground. NEW & NOTEWORTHY: Accurate foot placement is essential for safe walking. We used computational models and human walking experiments to test how our nervous system achieves this accuracy. We find that our control of foot placement beneficially combines sensory feedback with internal forward model predictions to accurately estimate the body's state. Our results match recent computational neuroscience findings for reaching movements, suggesting that state estimation is a general mechanism of human motor control.

Cutaneous reflex modulation during obstacle avoidance under conditions of normal and degraded visual input.

The nervous system integrates visual input regarding obstacles with limb-based sensory feedback to allow an individual to safely negotiate the environment. This latter source can include cutaneous information from the foot, particularly in the event that limb trajectory is not sufficient and there is an unintended collision with the object. However, it is not clear the extent to which cutaneous reflexes are modified based on visual input. In this study, we first determined if phase-dependent modulation of these reflexes is present when stepping over an obstacle during overground walking. We then tested the hypothesis that degrading the quality of visual feedback alters cutaneous reflex amplitude in this task. Subjects walked and stepped over an obstacle-leading with their right foot-while we electrically stimulated the right superficial peroneal nerve at the level of the ankle at different phases. Subjects performed this task with normal vision and with degraded vision. We found that the amplitude of cutaneous reflexes varied based on the phase of stepping over the obstacle in all leg muscles tested. With degraded visual feedback, regardless of phase, we found larger facilitation of cutaneous reflexes in the ipsilateral biceps femoris-a muscle responsible for flexing the knee to avoid the obstacle. Although degrading vision caused minor changes in several other muscles, none of these differences reached the level of significance. Nonetheless, our results suggest that visual feedback plays a role in altering how the nervous system uses other sensory input in a muscle-specific manner to ensure safe obstacle clearance.

Modification of cutaneous reflexes during visually guided walking.

Although it has become apparent that cutaneous reflexes can be adjusted based on the phase and context of the locomotor task, it is not clear to what extent these reflexes are regulated when locomotion is modified under visual guidance. To address this, we compared the amplitude of cutaneous reflexes while subjects performed walking tasks that required precise foot placement. In one experiment, subjects walked overground and across a horizontal ladder with narrow raised rungs. In another experiment, subjects walked and stepped onto a series of flat targets, which required different levels of precision (large vs. narrow targets). The superficial peroneal or tibial nerve was electrically stimulated in multiple phases of the gait cycle in each condition and experiment. Reflexes between 50 and 120 ms poststimulation were sorted into 10 equal phase bins, and the amplitudes were then averaged. In each experiment, differences in cutaneous reflexes between conditions occurred predominantly during swing phase when preparation for precise foot placement was necessary. For instance, large excitatory cutaneous reflexes in ipsilateral tibialis anterior were present in the ladder condition and when stepping on narrow targets compared with inhibitory responses in the other conditions, regardless of the nerve stimulated. In the ladder experiments, additional effects of walking condition were evident during stance phase when subjects had to balance on the narrow ladder rungs and may be related to threat and/or the unstable foot-surface interaction. Taken together, these results suggest that cutaneous reflexes are modified when visual feedback regarding the terrain is critical for successful walking.

Effects of age-related macular degeneration and ambient light on curb negotiation.

PURPOSE: To determine how age-related macular degeneration (AMD) and changes in ambient light affect the ability to negotiate a curb while walking. METHODS: Ten older adults with AMD and 11 normal-sighted control subjects performed a curb negotiation task under normal light (∼600 lux), dim light (∼0.7 lux), and following a sudden reduction (∼600 to 0.7 lux) of light. In this task, subjects walked and stepped up or down a simulated sidewalk curb. Movement kinematics and ground reaction forces were measured during curb ascent and descent. Habitual visual acuity, contrast sensitivity, and visual fields were also assessed. RESULTS: Apart from slower gait speed in those with AMD, there were no differences between groups during curb ascent for any other measure. During curb descent, older adults with AMD frequently used shuffling steps in the approach phase to locate the curb edge and showed prolonged double support duration stepping over the curb compared with control subjects. However, reduced lighting, particularly a sudden reduction, led to several significant changes in movement characteristics in both groups. For instance, toe clearance stepping up the curb was greater, and landing force stepping down was reduced. In addition, slower gait speed and greater double support duration were evident in curb ascent and descent. In AMD subjects, contrast sensitivity, visual acuity, and visual field threshold were associated with several kinematic measures in the three light conditions during curb negotiation. CONCLUSIONS: Minor AMD-specific changes in movement are seen during curb negotiation. However, attenuated lighting greatly impacts curb ascent and descent, regardless of eye disease, which manifests as a cautious walking strategy and may increase the risk of falling. Environmental enhancements that reduce the deleterious effects of poor lighting are required to improve mobility and quality of life of older adults, particularly those with AMD.

Effect of ambient light and age-related macular degeneration on precision walking.

PURPOSE: To determine how age-related macular degeneration (AMD) and changes in ambient light affect the control of foot placement while walking. METHODS: Ten older adults with AMD and 11 normal-sighted controls performed a precision walking task under normal (∼600 lx), dim (∼0.7 lx), and after a sudden reduction (∼600 to 0.7 lx) of light. The precision walking task involved subjects walking and stepping to the center of a series of irregularly spaced, low-contrast targets. Habitual visual acuity and contrast sensitivity and visual field function were also assessed. RESULTS: There were no differences between groups when performing the walking task in normal light (p > 0.05). In reduced lighting, older adults with AMD were less accurate and more variable when stepping across the targets compared to controls (p < 0.05). A sudden reduction of light proved the most challenging for this population. In the AMD group, contrast sensitivity and visual acuity were not significantly correlated with walking performance. Visual field thresholds in the AMD group were only associated with greater foot placement error and variability in the dim light walking condition (r = -0.69 to -0.87, p < 0.05). CONCLUSIONS: While walking performance is similar between groups in normal light, poor ambient lighting results in decreased foot placement accuracy in older adults with AMD. Improper foot placement while walking can lead to a fall and possible injury. Thus, to improve the mobility of those with AMD, strategies to enhance the environment in reduced lighting situations are necessary.

Changes in task parameters during walking prism adaptation influence the subsequent generalization pattern.

An understanding of the transfer (or generalization) of motor adaptations between legs and across tasks during walking has remained elusive due to limited research and mixed results. Here, we asked whether stepping sequences or task constraints introduced during walking prism-adaptation tasks influence generalization patterns. Forty subjects adapted to prism glasses in precision-walking or obstacle-avoidance tasks that required a specific stepping sequence to the center of two/three targets or laterally over an obstacle. We then tested for generalization, reflected by aftereffects in the nonadapted task. Our previous study using these tasks found that only one leg generalized. Here, we reversed the stepping sequence and found that only the opposite leg generalized in the subject group that adapted in a precision-walking task. The combination of stepping sequence and direction of prism shift caused subjects in two groups to collide with the obstacle early during adaptation, thus making the step prior to going over the obstacle more important. Both legs subsequently generalized. A fourth subject group experienced a three-target, precision-walking task, resulting in a balanced, right-left, left-right stepping sequence, meant to induce bilateral generalization. While only one leg generalized, foot placement aftereffects before stepping over the obstacle would have caused subjects to collide with it. Together with our previous study, the results suggest a contribution of stepping sequence during the adapted task on generalization patterns, likely driven by proprioceptive feedback. The results also support the idea that negative consequences during adaptation and/or perceived threat can influence generalization.

The effects of prior exposure to prism lenses on de novo motor skill learning.

Motor learning involves plasticity in a network of brain areas across the cortex and cerebellum. Such traces of learning have the potential to affect subsequent learning of other tasks. In some cases, prior learning can interfere with subsequent learning, but it may be possible to potentiate learning of one task with a prior task if they are sufficiently different. Because prism adaptation involves extensive neuroplasticity, we reasoned that the elevated excitability of neurons could increase their readiness to undergo structural changes, and in turn, create an optimal state for learning a subsequent task. We tested this idea, selecting two different forms of learning tasks, asking whether exposure to a sensorimotor adaptation task can improve subsequent de novo motor skill learning. Participants first learned a new visuomotor mapping induced by prism glasses in which prism strength varied trial-to-trial. Immediately after and the next day, we tested participants on a mirror tracing task, a form of de novo skill learning. Prism-trained and control participants both learned the mirror tracing task, with similar reductions in error and increases in distance traced. Both groups also showed evidence of offline performance gains between the end of day 1 and the start of day 2. However, we did not detect differences between groups. Overall, our results do not support the idea that prism adaptation learning can potentiate subsequent de novo learning. We discuss factors that may have contributed to this result.

The contribution of vision, proprioception, and efference copy in storing a neural representation for guiding trail leg trajectory over an obstacle.

Stepping over obstacles requires vision to guide the leading leg, but direct visual information is not available to guide the trailing leg. The neural mechanisms for establishing a stored obstacle representation and thus facilitating the trail leg trajectory in humans are unknown. Twenty-four subjects participated in one of three experiments, which were designed to investigate the contribution of visual, proprioceptive, and efference copy signals. Subjects stepped over an obstacle with their lead leg, stopped, and straddled the obstacle for a delay period before stepping over it with their trail leg while toe elevation was recorded. Subsequently, we calculated maximum toe elevation and toe clearance. First, we found that subjects could accurately scale trail leg toe elevation and clearance, despite straddling an obstacle for up to 2 min, similar to quadrupeds. Second, we found that when the lead leg was passively moved over an obstacle (eliminating an efference copy signal and altering proprioception) without vision, trail leg toe elevation and clearance were reduced, and variability increased compared with when subjects actively moved their lead leg. Trail leg toe measures returned to normal when vision was provided during the passive manipulation. Finally, we found that altering lead leg proprioceptive feedback by adding mass to the ankle had no effect on trail leg toe measures. Taken together, our results suggest that humans can store a neural representation of obstacle properties for extended periods of time and that vision appears to be sufficient in this process to guide trail leg trajectory.

Learning from the Physical Consequences of Our Actions Improves Motor Memory.

Actions have consequences. Motor learning involves correcting actions that lead to movement errors and remembering these actions for future behavior. In most laboratory situations, movement errors have no physical consequences and simply indicate the progress of learning. Here, we asked how experiencing a physical consequence when making a movement error affects motor learning. Two groups of participants adapted to a new, prism-induced mapping between visual input and motor output while performing a precision walking task. Importantly, one group experienced an unexpected slip perturbation when making foot-placement errors during adaptation. Because of our innate drive for safety, and the fact that balance is fundamental to movement, we hypothesized that this experience would enhance motor memory. Learning generalized to different walking tasks to a greater extent in the group who experienced the adverse physical consequence. This group also showed faster relearning one week later despite exposure to a competing mapping during initial learning, evidence of greater memory consolidation. The group differences in generalization and consolidation occurred although they both experienced similar magnitude foot-placement errors and adapted at similar rates. Our results suggest the brain considers the potential physical consequences of movement error when learning and that balance-threatening consequences serve to enhance this process.

Probing the deployment of peripheral visual attention during obstacle-crossing planning.

Gaze is directed to one location at a time, making peripheral visual input important for planning how to negotiate different terrain during walking. Whether and how the brain attends to this input is unclear. We developed a novel paradigm to probe the deployment of sustained covert visual attention by testing orientation discrimination of a Gabor patch at stepping and non-stepping locations during obstacle-crossing planning. Compared to remaining stationary, obstacle-crossing planning decreased visual performance (percent correct) and sensitivity (d') at only the first of two stepping locations. Given the timing of the first and second steps before obstacle crossing relative to the Gabor patch presentation, the results suggest the brain uses peripheral vision to plan one step at a time during obstacle crossing, in contrast to how it uses central vision to plan two or more steps in advance. We propose that this protocol, along with multiple possible variations, presents a novel behavioral approach to identify the role of covert visual attention during obstacle-crossing planning and other goal-directed walking tasks.

Contribution of cells in the posterior parietal cortex to the planning of visually guided locomotion in the cat: effects of temporary visual interruption.

In the present study, we determined whether cells in the posterior parietal cortex (PPC) may contribute to the planning of voluntary gait modifications in the absence of visual input. In two cats we recorded the responses of 41 neurons in layer V of the PPC that discharged in advance of the gait modification to a 900-ms interruption of visual information (visual occlusion). The cats continued to walk without interruption during the occlusion, which produced only minimal changes in step cycle duration and paw placement. Visual occlusion applied during the period of cell discharge was without significant effect on discharge frequency in 57% of cells. In the other cells, the visual occlusion produced either significant decreases (18%) or increases (21%) of discharge activity (in 1 cell there was both an increase and a decrease). The mean latency of the changes was 356 ms for decreases and 252 ms for increases. In most neurons, discharge frequency, when modified, returned to the same levels as during unoccluded locomotion when vision was restored. In some cells, there were significant changes in discharge activity after the restoration of vision; these were associated with corrections of gait. These results suggest that the PPC is more involved in the visuomotor transformations necessary to plan gait modifications than in continual sensory processing of visual information. We further propose that cells in the PPC contribute both to the planning of gait modifications on the basis of only intermittent visual sampling and to visually guided online corrections of gait.