Response preparation of a secondary reaction time task is influenced by movement phase within a continuous visuomotor tracking task.

The simultaneous performance of two motor tasks is challenging. Currently, it is unclear how response preparation of a secondary task is impacted by the performance of a continuous primary task. The purpose of the present experiment was to investigate whether the position of the limb performing the primary cyclical tracking task impacts response preparation of a secondary reaction time task. Participants (n = 20) performed a continuous tracking task with their left hand that involved cyclical and targeted wrist flexion and extension. Occasionally, a probe reaction time task requiring isometric wrist extension was performed with the right hand in response to an auditory stimulus (80 or 120 dB) that was triggered when the left hand passed through one of 10 locations identified within the movement cycle. On separate trials, transcranial magnetic stimulation was applied over the left primary motor cortex and triggered at the same 10 stimulus locations to assess corticospinal excitability associated with the probe reaction time task. Results revealed that probe reaction times were significantly longer and motor-evoked potential amplitudes were significantly larger when the left hand was in the middle of a movement cycle compared with an endpoint, suggesting that response preparation of a secondary probe reaction time task was modulated by the phase of movement within the continuous primary task. These results indicate that primary motor task requirements can impact preparation of a secondary task, reinforcing the importance of considering primary task characteristics in dual-task experimental design.

Improved proprioception does not benefit visuomotor adaptation.

Visuomotor adaptation arises when reaching in an altered visual environment, where one's seen hand position does not match their felt (i.e., proprioceptive) hand position in space. Here, we asked if proprioceptive training benefits visuomotor adaptation, and if these benefits arise due to implicit (unconscious) or explicit (conscious strategy) processes. Seventy-two participants were divided equally into 3 groups: proprioceptive training with feedback (PTWF), proprioceptive training no feedback (PTNF), and Control (CTRL). The PTWF and PTNF groups completed passive proprioceptive training, where a participant's hand was moved to an unknown reference location and they judged the felt position of their unseen hand relative to their body midline on every trial. The PTWF group received verbal feedback with respect to their response accuracy on the middle 60% of trials, whereas the PTNF did not receive any feedback during training. The CTRL group did not complete proprioceptive training and instead sat quietly during this time. Following proprioceptive training or time delay, all three groups reached when seeing a cursor that was rotated 30° clockwise relative to their hand motion. The experiment ended with participants completing a series of no-cursor reaches to assess implicit and explicit adaptation. Results indicated that the PTWF group improved the accuracy of their sense of felt hand position following proprioceptive training. However, this improved proprioceptive acuity (i.e., the accuracy of their sense of felt hand) did not benefit visuomotor adaptation, as all three groups showed similar visuomotor adaptation across rotated reach training trials. Visuomotor adaptation arose implicitly, with minimal explicit contribution for all three groups. Together, these results suggest that passive proprioceptive training does not benefit, nor hinder, the extent of implicit visuomotor adaptation established immediately following reach training with a 30° cursor rotation.

The influence of awareness on implicit visuomotor adaptation.

It is well documented that reaches are adapted when reaching with a visuomotor distortion (i.e., rotated cursor feedback). Less clear is the influence of awareness on visuomotor adaptation, where awareness encompasses knowledge of the changes in one's reaches and the visuomotor distortion itself. In the current experiment, we asked if awareness governs the magnitude of implicit (i.e., unconscious) visuomotor adaptation achieved, independent of how the distortion is introduced (i.e., abruptly vs. gradually introduced visuomotor distortion), and hence initial errors experienced. Participants were divided into two groups that differed with respect to how the visuomotor distortion was introduced (i.e., Abrupt vs. Gradual Groups) and reached in a virtual environment where a cursor on the screen misrepresented the position of their hand. Participants completed three blocks of 150 reach training trials in the following order: aligned cursor feedback (baseline), rotated cursor feedback (adaptation) and aligned cursor feedback (washout). For the Abrupt Group, the cursor was immediately rotated 45° clockwise (CW) relative to hand motion in the adaptation block, whereas in the Gradual Group, the 45° cursor rotation was gradually introduced over adaptation trials. Following reach training, participants' awareness of changes in their reaches and the visuomotor distortion were established based on a drawing task, where participants drew the path their hand took to get the cursor on target, as well as a post-experiment questionnaire. Participants were subsequently divided into the following 3 groups: Abrupt-Aware (n = 16), Gradual-Aware (n = 11) and Gradual-Unaware (n = 14). Results revealed that errors differed for the Gradual-Unaware Group at the end of adaptation training compared to the Gradual-Aware Group and at the start of the washout block compared to the Abrupt-Aware Group. Errors in the two aware groups did not differ from each other. These results suggest that awareness may lead to reduced implicit adaptation, regardless of the size of initial errors experienced.

Age-related changes in upper limb coordination in a complex reaching task.

When reaching to targets within arm's reach, intentional trunk motion must be neutralized by compensatory motion of the upper limb (UL). Advanced age has been associated with deterioration in the coordination of multi-joint UL movements. In the current study, we looked to determine if older adults also have difficulties modifying their UL movements (i.e., coordination between the shoulder and elbow joints), during a complex reaching task when trunk motion is manipulated. Two groups of healthy participants were recruited: 18 young (mean age = 24.28 ± 2.89 years old) and 18 older (mean age = 72.11 ± 2.39 years old) adults. Participants reached to a target with their eyes closed, while simultaneously moving the trunk forward. In 40% of trials, the trunk motion was unexpectedly blocked. Participants performed the task with both their dominant and non-dominant arms, and at a preferred and fast speed. All participants were able to coordinate motion at the elbow and shoulder joints in a similar manner and modify this coordination in accordance with motion at the trunk, regardless of the hand used or speed of movement. Specifically, in reaches that involved forward trunk motion (free-trunk trials), all participants demonstrated increased elbow flexion (i.e., less elbow extension) compared to blocked-trunk trials. In contrast, when trunk motion was blocked (blocked-trunk trials), all reaching movements were accompanied by increased shoulder horizontal adduction. While coordination of UL joints was similar across older and young adults, the extent of changes at the elbow and shoulder was smaller and less consistent in older adults compared to young participants, especially when trunk motion was involved. These results suggest that older adults can coordinate their UL movements based on task requirements, but that their performance is not as consistent as young adults.

Proprioceptive recalibration following implicit visuomotor adaptation is preserved in Parkinson's disease.

Individuals with Parkinson's disease (PD) and healthy adults demonstrate similar levels of visuomotor adaptation provided that the distortion is small or introduced gradually, and hence, implicit processes are engaged. Recently, implicit processes underlying visuomotor adaptation in healthy individuals have been proposed to include proprioceptive recalibration (i.e., shifts in one's proprioceptive sense of felt hand position to match the visual estimate of their hand experienced during reaches with altered visual feedback of the hand). In the current study, we asked if proprioceptive recalibration is preserved in PD patients. PD patients tested during their "off" and "on" medication states and age-matched healthy controls reached to visual targets, while visual feedback of their unseen hand was gradually rotated 30° clockwise or translated 4 cm rightwards of their actual hand trajectory. As expected, PD patients and controls produced significant reach aftereffects, indicating visuomotor adaptation after reaching with the gradually introduced visuomotor distortions. More importantly, following visuomotor adaptation, both patients and controls showed recalibration in hand position estimates, and the magnitude of this recalibration was comparable between PD patients and controls. No differences for any measures assessed were observed across medication status (i.e., PD off vs PD on). Results reveal that patients are able to adjust their sensorimotor mappings and recalibrate proprioception following adaptation to a gradually introduced visuomotor distortion, and that dopaminergic intervention does not affect this proprioceptive recalibration. These results suggest that proprioceptive recalibration does not involve striatal dopaminergic pathways and may contribute to the preserved visuomotor adaptation that arises implicitly in PD patients.

Going offline: differences in the contributions of movement control processes when reaching in a typical versus novel environment.

Human movements are remarkably adaptive. We are capable of completing movements in a novel visuomotor environment with similar accuracy to those performed in a typical environment. In the current study, we examined if the control processes underlying movements under typical conditions were different from those underlying novel visuomotor conditions. 16 participants were divided into two groups, one receiving continuous visual feedback during all reaches (CF), and the other receiving terminal feedback regarding movement endpoint (TF). Participants trained in a virtual environment by completing 150 reaches to three targets when (1) a cursor accurately represented their hand motion (i.e., typical environment) and (2) a cursor was rotated 45° clockwise relative to their hand motion (i.e., novel environment). Analyses of within-trial measures across 150 reaching trials revealed that participants were able to demonstrate similar movement outcomes (i.e., movement time and angular errors) regardless of visual feedback or reaching environment by the end of reach training. Furthermore, a reduction in variability across several measures (i.e., reaction time, movement time, time after peak velocity, and jerk score) over time showed that participants improved the consistency of their movements in both reaching environments. However, participants took more time and were less consistent in the timing of initiating their movements when reaching in a novel environment compared to reaching in a typical environment, even at the end of training. As well, angular error variability at different proportions of the movement trajectory was consistently greater when reaching in a novel environment across trials and within a trial. Together, the results suggest a greater contribution of offline control processes and less effective online corrective processes when reaching in a novel environment compared to when reaching in a typical environment.

Age differences in arm-trunk coordination during trunk-assisted reaching.

Reaching for an object is a basic motor skill that requires precise coordination between elbow, shoulder and trunk motion. The purpose of this research study was to examine age-related differences in compensatory arm-trunk coordination during trunk-assisted reaching. To engage the arm and trunk, an older and younger group of participants were asked to (1) maintain a fixed hand position while flexing forward at the trunk [stationary hand task (SHT)] and (2) reach to a within-arm's reach target while simultaneously flexing forward at the trunk [reaching hand task (RHT)] (Raptis et al. in J Neurophysiol 97:4069-4078, 2007; Sibindi et al. in J Vestib Res 23:237-247, 2013). Both tasks were completed with eyes closed. Participants completed the two tasks with their dominant and non-dominant arms, and at both a fast and a preferred speed. On average, young and older participants performed in a similar manner in the SHT, such that they maintained their hand position by compensating for trunk movement with modifications of the elbow and shoulder joints. In the RHT, young and older participants had similar endpoint accuracy. This similarity in performance between young and older participants in the SHT and RHT tasks was observed regardless of the arm used or movement speed. However, for both tasks, movements in older adults were significantly more variable compared to younger adults as shown by the larger variability in arm-trunk coordination performance (gain scores) in the SHT and higher movement time variability in the RHT. Thus, results imply that older adults maintain their ability to coordinate arm and trunk movements efficiently during reaching actions but are not as consistent as younger adults.

The influence of awareness on explicit and implicit contributions to visuomotor adaptation over time.

Explicit (strategic) and implicit (unconscious) processes play a role in visuomotor adaptation (Bond and Taylor, J Neurophysiol 113:3836-3849, https://doi.org/10.1152/jn.00009.2015 , 2015; Werner et al., PLoS ONE 10:1-18, https://doi.org/10.1371/journal.pone.0123321 , 2015). We investigated the contributions of explicit and implicit processes to visuomotor adaptation when awareness was manipulated directly vs. indirectly, and asked how these contributions changed over time. Participants were assigned to a Strategy or No-Strategy group. Those in the Strategy group were made aware of the visuomotor distortion directly. Participants were further subdivided into groups to train with a large (60°), medium (40°) or small (20°) visuomotor distortion, providing the potential for awareness to develop indirectly. Participants reached with their respective distorted cursor, followed by a series of no-cursor reaches to assess the contributions of explicit and implicit processes to visuomotor adaptation after every 30 reach training trials. Within the no-cursor reaching trials, participants reached (1) with any strategies they had gained during training (explicit + implicit processes), and (2) as accurately to the target as possible (implicit processes). Results showed that implicit contributions were greatest in the No-Strategy group, took time to develop, and were transient, as partial decay was seen following a 5-min rest. As well, implicit contributions were similar (i.e., plateaued), regardless of the rotation size participants trained with. In contrast, explicit contributions were greatest in the Strategy group, increased with rotation size, and remained consistent over time. Taken together, results reveal that there are notable differences in the stability of explicit and implicit processes and their potential to contribute to visuomotor adaptation depending on if awareness is provided directly.

Adaptation to proprioceptive targets following visuomotor adaptation.

In the following study, we asked if reaches to proprioceptive targets are updated following reach training with a gradually introduced visuomotor perturbation. Subjects trained to reach with distorted hand-cursor feedback, such that they saw a cursor that was rotated or translated relative to their actual hand movement. Following reach training trials with the cursor, subjects reached to Visual (V), Proprioceptive (P) and Visual + Proprioceptive (VP) targets with no visual feedback of their hand. Comparison of reach endpoints revealed that reaches to VP targets followed similar trends as reaches to P targets, regardless of the training distortion introduced. After reaching with a rotated cursor, subjects adapted their reaches to all target types in a similar manner. However, after reaching with a translated cursor, subjects adapted their reach to V targets only. Taken together, these results show that following training with a visuomotor distortion, subjects primarily rely on proprioceptive information when reaching to VP targets. Furthermore, results indicate that reach adaptation to P targets depends on the distortion presented. Training with a rotation distortion leads to changes in reaches to both V and P targets, while a translation distortion, which introduces a constant discrepancy between visual and proprioceptive estimates of hand position throughout the reach, affects changes to V but not P targets.

Go-activation endures following the presentation of a stop-signal: evidence from startle.

It has been proposed that, in a stop-signal task (SST), independent go- and stop-processes "race" to control behavior. If the go-process wins, an overt response is produced, whereas, if the stop-process wins, the response is withheld. One prediction that follows from this proposal is that, if the activation associated with one process is enhanced, it is more likely to win the race. We looked to determine whether these initiation and inhibition processes (and thus response outcomes) could be manipulated by using a startling acoustic stimulus (SAS), which has been shown to provide additional response activation. In the present study, participants were to respond to a visual go-stimulus; however, if a subsequent stop-signal appeared, they were to inhibit the response. The stop-signal was presented at a delay corresponding to a probability of responding of 0.4 (determined from a baseline block of trials). On stop-trials, a SAS was presented either simultaneously with the go-signal or stop-signal or 100, 150, or 200 ms following the stop-signal. Results showed that presenting a SAS during stop-trials led to an increase in probability of responding when presented with or following the stop-signal. The latency of SAS responses at the stop-signal + 150 ms and stop-signal + 200 ms probe times suggests that they would have been voluntarily inhibited but instead were involuntarily initiated by the SAS. Thus results demonstrate that go-activation endures even 200 ms following a stop-signal and remains accessible well after the response has been inhibited, providing evidence against a winner-take-all race between independent go- and stop-processes. NEW & NOTEWORTHY: In this study, a startling acoustic stimulus (SAS) was used to determine whether response outcome could be manipulated in a stop-signal task. Results revealed that presenting a SAS during stop-signal trials led to an increase in probability of responding even when presented 200 ms following the stop-signal. The latency of SAS responses indicates that go-activation remains accessible and modifiable well after the response is voluntarily inhibited, providing evidence against an irrevocable commitment to inhibition.

Startle reveals decreased response preparatory activation during a stop-signal task.

In a stop-signal task participants are instructed to initiate a movement in response to a go signal, but to inhibit this movement if an infrequent stop signal is presented after the go. Reaction time (RT) in a stop-signal task is typically longer compared with that in a simple RT task, which may be attributed to a reduced readiness to initiate the response caused by the possibility of having to inhibit the response. The purpose of this experiment was to probe the preparatory activation level of the motor response during a stop-signal task using a startling acoustic stimulus (SAS), which has been shown to involuntarily trigger sufficiently prepared responses at a short latency. Participants completed two separate tasks: a simple RT task, followed by a stop-signal RT task. During both tasks, an SAS (120 dB) was pseudorandomly presented concurrently with the go signal. As expected, RT during the simple RT task was significantly shorter than during the stop-signal task. A significant reduction in RT was noted when an SAS was presented during the simple RT task; however, during the stop-signal task, an SAS resulted in either a significant speeding or a moderate delay in RT. Additionally, the subset of SAS trial responses with the shortest RT latencies produced during the stop-signal task were also delayed compared with the short-latency SAS trial responses observed during the simple RT task. Despite evidence that a response was prepared in advance of the go signal during a stop-signal task, it appears that the amount of preparatory activation was reduced compared with that achieved during a simple RT task.

Proprioceptive recalibration arises slowly compared to reach adaptation.

When subjects reach in a novel visuomotor environment (e.g. while viewing a cursor representing their hand that is rotated from their hand's actual position), they typically adjust their movements (i.e. bring the cursor to the target), thus reducing reaching errors. Additionally, research has shown that reaching with altered visual feedback of the hand results in sensory changes, such that proprioceptive estimates of hand position are shifted in the direction of the visual feedback experienced (Cressman and Henriques in J Neurophysiol 102:3505-3518, 2009). This study looked to establish the time course of these sensory changes. Additionally, the time courses of implicit sensory and motor changes were compared. Subjects reached to a single visual target while seeing a cursor that was either aligned with their hand position (50 trials) or rotated 30° clockwise relative to their hand (150 trials). Reach errors and proprioceptive estimates of felt hand position were assessed following the aligned reach training trials and at seven different times during the rotated reach training trials by having subjects reach to the target without visual feedback, and provide estimates of their hand relative to a visual reference marker, respectively. Results revealed a shift in proprioceptive estimates throughout the rotated reach training trials; however, significant sensory changes were not observed until after 70 trials. In contrast, results showed a greater change in reaches after a limited number of reach training trials with the rotated cursor. These findings suggest that proprioceptive recalibration arises more slowly than reach adaptation.

The rapid-chase theory does not extend to movement execution.

It is assumed that the processing of a prime followed by a mask occurs sequentially in a feedforward manner when the three (initiation, takeover, and independence) criteria outlined by the rapid-chase theory are met. The purpose of the current study was to determine if the processing of the prime and mask fit the predictions of the rapid-chase theory when the prime and mask are presented during an ongoing movement. In two experiments, participants made rapid pointing movements to a target indicated by the mask. In Experiment 1, the prime was presented at movement onset and the prime-mask stimulus onset asynchrony (SOA) was manipulated. In Experiment 2, the prime-mask SOA was constant but the delay between movement and prime onset was manipulated. Although the results support the initiation and takeover criteria, the data did not support the independence criterion. Consequently, the rapid-chase theory does not appear to extend to movement execution.

Generalization patterns for reach adaptation and proprioceptive recalibration differ after visuomotor learning.

Visuomotor learning results in changes in both motor and sensory systems (Cressman EK, Henriques DY. J Neurophysiol 102: 3505-3518, 2009), such that reaches are adapted and sense of felt hand position recalibrated after reaching with altered visual feedback of the hand. Moreover, visuomotor learning has been shown to generalize such that reach adaptation achieved at a trained target location can influence reaches to novel target directions (Krakauer JW, Pine ZM, Ghilardi MF, Ghez C. J Neurosci 20: 8916-8924, 2000). We looked to determine whether proprioceptive recalibration also generalizes to novel locations. Moreover, we looked to establish the relationship between reach adaptation and changes in sense of felt hand position by determining whether proprioceptive recalibration generalizes to novel targets in a similar manner as reach adaptation. On training trials, subjects reached to a single target with aligned or misaligned cursor-hand feedback, in which the cursor was either rotated or scaled in extent relative to hand movement. After reach training, subjects reached to the training target and novel targets (including targets from a second start position) without visual feedback to assess generalization of reach adaptation. Subjects then performed a proprioceptive estimation task, in which they indicated the position of their hand relative to visual reference markers placed at similar locations as the trained and novel reach targets. Results indicated that shifts in hand position generalized across novel locations, independent of reach adaptation. Thus these distinct sensory and motor generalization patterns suggest that reach adaptation and proprioceptive recalibration arise from independent error signals and that changes in one system cannot guide adjustments in the other.

Retention of proprioceptive recalibration following visuomotor adaptation.

We have recently shown that visuomotor adaptation following reaches with a misaligned cursor not only induces changes in an individual's motor output, but their proprioceptive sense of hand position as well. Long-term changes are seen in motor adaptation; however, very little is known about the retention of changes in felt hand position. We sought to evaluate whether this recalibration in proprioception, following visuomotor adaptation, is sufficiently robust to be retained the following day (~24 h later), and if so, to determine its extent. Visuomotor adaptation was induced by having subjects perform reaches to visual targets using a cursor representing their unseen hand, which had been gradually rotated 45° counterclockwise. Motor adaptation and proprioceptive recalibration were determined by assessing subjects' reach aftereffects and changes in hand bias, respectively. We found that subjects adapted their reaches and recalibrated their sense of hand position following training with a misaligned cursor, as shown in Cressman and Henriques (J Neurophysiol 102:3505-3518, 2009). More importantly, subjects who showed proprioceptive recalibration in the direction of motor adaptation on Day 1 did retain changes in felt hand position and motor adaptation on Day 2. These findings suggest that in addition to motor changes, individuals are capable of retaining sensory changes in proprioception up to 24 h later.

Generalization of reach adaptation and proprioceptive recalibration at different distances in the workspace.

Studies have shown that adapting one's reaches in one location in the workspace can generalize to other novel locations. Generalization of this visuomotor adaptation is influenced by the location of novel targets relative to the trained location such that reaches made to novel targets that are located far from the trained target direction (i.e., ~22.5°; Krakauer et al. in J Neurosci 20:8916-8924, 2000) show very little generalization compared to those that are closer to the trained direction. However, generalization is much broader when reaching to novel targets in the same direction but at different distances from the trained target. In this study, we investigated whether changes in hand proprioception (proprioceptive recalibration), like reach adaptation, generalize to different distances of the workspace. Subjects adapted their reaches with a rotated cursor to two target locations at a distance of 13 cm from the home position. We then compared changes in open-loop reaches and felt hand position at these trained locations to novel targets located in the same direction as the trained targets but either at a closer (10 cm) or at a farther distance (15 cm) from the home position. We found reach adaptation generalized to novel closer and farther targets to the same extent as observed at the trained target distance. In contrast, while changes in felt hand position were significant across the two novel distances, this recalibration was smaller for the novel-far locations compared to the trained location. Given that reach adaptation completely generalized across the novel distances but proprioceptive recalibration generalized to a lesser extent for farther distances, we suggest that proprioceptive recalibration may arise independently of motor adaptation and vice versa.

Sensory integration during reaching: the effects of manipulating visual target availability.

When using visual and proprioceptive information to plan a reach, it has been proposed that the brain combines these cues to estimate the object and/or limb's location. Specifically, according to the maximum-likelihood estimation (MLE) model, sensory inputs are combined such that more reliable inputs are assigned a greater weight (Ernst and Banks in Nature 415:429-433, 2002). In this paper, we examined if the brain is able to adjust which sensory cue it weights the most. Specifically, we asked if the brain changes how it weights sensory information when the availability of a visual cue is manipulated. Twelve healthy subjects reached to visual (V), proprioceptive (P), or visual + proprioceptive (VP) targets under different visual delay conditions (e.g., on V and VP trials, the visual target was available for the entire reach; it was removed with the go signal, or it was removed 1 s before the go signal). To establish which sensory cue subjects weighted the most, we compared endpoint positions achieved on V and P reaches to VP reaches. Results indicated that subjects combined visual and proprioceptive cues in accordance with the MLE model when reaching to VP targets. Moreover, subjects' reaching errors to visual targets increased with longer visual delays (particularly in the vertical direction). However, there was no change in reach variability with longer delays, and subjects did not reweight visual information as the availability of visual information was manipulated. Thus, a change in visual environment is not sufficient to cause the brain to reweight how it processes sensory information.

The effect of visuomotor adaptation on proprioceptive localization: the contributions of perceptual and motor changes.

Reaching movements are rapidly adapted following training with rotated visual feedback of the hand (motor recalibration). Our laboratory has also found that visuomotor adaptation results in changes in estimates of felt hand position (proprioceptive recalibration) in the direction of the visuomotor distortion (Cressman and Henriques 2009, 2010; Cressman et al. 2010). In the present study, we included an additional method for measuring hand proprioception [specifically, proprioceptive-guided reaches of the unadapted (left) hand to the robot-guided adapted (right) hand-target] and compared this with our original perceptual task (estimating the felt hand position of the adapted hand relative to visual reference markers/the body midline), as well as to no-cursor reaches with the adapted hand (reaching to visual and midline-targets), to better identify whether changes in reaching following adaptation to a 50° rightward-rotated cursor reflect sensory or motor processes. Results for the proprioceptive estimation task were consistent with previous findings; subjects felt their hand to be aligned with a reference marker when it was shifted approximately 4° more in the direction of the visuomotor distortion following adaptation compared with baseline conditions. Moreover, we found similar changes in the proprioceptive-guided reaching task such that subjects misreached 5° in the direction of the cursor rotation. However, these results were true only for proprioceptive-guided reaches to the adapted hand, as reaches to the body midline were not affected by adaptation. This suggests that proprioceptive recalibration is restricted to the adapted hand and does not generalize to the rest of the body; this truly reflects a change in the sensory representation of the hand rather than changes in the motor program. This is in contrast to no-cursor reaches made with the adapted hand, which show reach after-effects for both visual targets and the midline, suggesting that reaches with the adapted hand reflect more of a change in the motor system. Our results also shed light on previous studies that may have misattributed these sensory and motor changes.

Intermanual transfer and proprioceptive recalibration following training with translated visual feedback of the hand.

Reaching with visual feedback that is misaligned with respect to the actual hand's location leads to changes in reach trajectories (i.e., visuomotor adaptation). Previous studies have also demonstrated that when training to reach with misaligned visual feedback of the hand, the opposite hand also partially adapts, providing evidence of intermanual transfer. Moreover, our laboratory has shown that visuomotor adaptation to a misaligned hand cursor, either translated or rotated relative to the hand, also leads to changes in felt hand position (what we call proprioceptive recalibration), such that subjects' estimate of felt hand position relative to both visual and non-visual reference markers (e.g., body midline) shifts in the direction of the visuomotor distortion. In the present study, we first determined the extent that motor adaptation to a translated cursor leads to transfer to the opposite hand, and whether this transfer differs across the dominant and non-dominant hands. Second, we looked to establish whether changes in hand proprioception that occur with the trained hand following adaptation also transfer to the untrained hand. We found intermanual motor transfer to the left untrained (non-dominant) hand after subjects trained their right (dominant) hand to reach with translated visual feedback of their hand. Motor transfer from the left trained to the right untrained hand was not observed. Despite finding changes in felt hand position in both trained hands, we did not find similar evidence of proprioceptive recalibration in the right or left untrained hands. Taken together, our results suggest that unlike visuomotor adaptation, proprioceptive recalibration does not transfer between hands and is specific only to the arm exposed to the distortion.

Comparing movement preparation of unimanual, bimanual symmetric, and bimanual asymmetric movements.

The goal of this study was to determine the process or processes most likely to be involved in reaction-time costs for spatially cued bimanual reaching. We used reaction time to measure the cost of bimanual symmetric movements compared to unimanual movements (a bimanual symmetric cost) and the cost for bimanual asymmetric movements compared to symmetric movements (a bimanual asymmetric cost). The results showed that reaction times were comparable for all types of movements in simple reaction time; that is, there was neither a bimanual symmetric cost nor an asymmetric cost. Therefore, unimanual, bimanual symmetric, and bimanual asymmetric movements have comparable complexity during response initiation. In choice conditions, there was no bimanual symmetric cost but there was a bimanual asymmetric cost, indicating that the preparation of asymmetric movements is more complex than symmetric movements. This asymmetric cost is likely the result of interference during response programming.