Patrick Bédard

On a basal ganglia role in learning and rehearsing visual-motor associations.

Fronto-striatal circuitry interacts with the midbrain dopaminergic system to mediate the learning of stimulus-response associations, and these associations often guide everyday actions, but the precise role of these circuits in forming and consolidating rules remains uncertain. A means to examine basal ganglia circuit contributions to associative motor learning is to examine these process in a lesion model system, such as Parkinsons disease (PD), a basal ganglia disorder characterized by the loss of dopamine neurons. We used functional magnetic resonance imaging (MRI) to compare brain activation of PD patients with a group of healthy aged-match participants during a visual-motor associative learning task that entailed discovering and learning arbitrary associations between a set of six visual stimuli and corresponding spatial locations by moving a joystick-controlled cursor. We tested the hypothesis that PD would recruit more areas than age-matched controls during learning and also show increased activation in commonly activated regions, probably in the parietal and premotor cortices, and the cerebellum, perhaps as compensatory mechanisms for their disrupted fronto-striatal networks. PD had no effect in acquiring the associative relationships and learning-related activation in several key frontal cortical and subcortical structures. However, we found that PD modified activation in other areas, including those in the cerebellum and frontal, and parietal cortex, particularly during initial learning. These results may suggest that the basal ganglia circuits become active more so during the initial formation of rule-based behavior.

On the role of static and dynamic visual afferent information in goal-directed aiming movements

Abstract. Movement planning has been shown to be optimized when the participant is permitted to see his or her hand resting on the starting base prior to movement initiation. However, this proposition is opposed by contradictory results. In the present study, we wanted to determine whether these conflicting results were caused by procedural differences. The results showed that seeing ones hand on the starting base did not result in more accurate aiming movement than when this information was not available. However, lower aiming errors were found when one was asked to foveate the starting base and then the target prior to movement initiation, but only when no dynamic visual information was available during movement. When an aiming movement was performed while ones hand was visible in visual periphery, foveating the starting base or not prior to movement initiation did not modify aiming accuracy. These results suggest that gazing at the starting base and then at the target provides an eye-based representation of the movement to be performed that can be used by the CNS to plan a manual aiming movement. Information for better planning of the direction – but not the extent – dimension of an upcoming movement can also be derived from dynamic visual information available in peripheral vision.

Gaze and Hand Position Effects on Finger-Movement-Related Human Brain Activation

Humans commonly use their hands to move and to interact with their environment by processing visual and proprioceptive information to determine the location of a goal-object and the initial hand position. It remains elusive, however, how the human brain fully uses this sensory information to generate accurate movements. In monkeys, it appears that frontal and parietal areas use and combine gaze and hand signals to generate movements, whereas in humans, prior work has separately assessed how the brain uses these two signals. Here we investigated whether and how the human brain integrates gaze orientation and hand position during simple visually triggered finger tapping. We hypothesized that parietal, frontal, and subcortical regions involved in movement production would also exhibit modulation of movement-related activation as a function of gaze and hand positions. We used functional MRI to measure brain activation while healthy young adults performed a visually cued finger movement and fixed gaze at each of three locations and held the arm in two different configurations. We found several areas that exhibited activation related to a mixture of these hand and gaze positions; these included the sensory-motor cortex, supramarginal gyrus, superior parietal lobule, superior frontal gyrus, anterior cingulate, and left cerebellum. We also found regions within the left insula, left cuneus, left midcingulate gyrus, left putamen, and right tempo-occipital junction with activation driven only by gaze orientation. Finally, clusters with hand position effects were found in the cerebellum bilaterally. Our results indicate that these areas integrate at least two signals to perform visual-motor actions and that these could be used to subserve sensory-motor transformations.

Movement planning of video and of manual aiming movements

We studied aiming performance of adults for video- and manual aiming tasks when they had visual information about the location of the starting base or when they had not. In video-aiming, foveating the starting base and then the target prior to movement initiation (Foveation) resulted in less aiming bias and variability than when the starting base was not visible (PNV), or visible without the participants foveating it prior to movement initiation (PSV). In manual aiming, Foveation and PSV procedures resulted in identical results but reduced aiming bias and variability in comparison to the PNV procedures. The results indicate that participants had difficulty in transforming the locations of the starting base and of the target when seen on a vertical screen into an appropriate movement trajectory. Successive foveation of the starting base and of the target facilitated this transformation, resulting in direction variability being reduced by more than half in comparison to the PNV and PSV conditions. This suggests that in video-aiming the efference copy of the saccade can be used by the CNS to approximate the hand trajectory in the workspace and/or in joint coordinates (Jouffrais and Boussaoud, 1999). Hand trajectory could be readily available in manual aiming if the target location can be recoded directly in hand-coordinates as recently suggested by Buneo et al. (2002).

On the role of peripheral visual afferent information for the control of rapid video-aiming movements

It has been shown that, even for very fast and short duration movements, seeing ones hand in peripheral vision, or a cursor representing it on a video screen, resulted in a better direction accuracy of a manual aiming movement than when the task was performed while only the target was visible. However, it is still unclear whether this was caused by on-line or off-line processes. Through a novel series of analyses, the goal of the present study was to shed some light on this issue. We replicated previous results showing that the visual information concerning ones movement, which is available between 40 degrees and 25 degrees of visual angle, is not useful to ensure direction accuracy of video-aiming movements, whereas visual afferent information available between 40 degrees and 15 degrees of visual angle improved direction accuracy over a target-only condition. In addition, endpoint variability on the direction component of the task was scaled to direction variability observed at peak movement velocity. Similar observations were made in a second experiment when the position of the cursor was translated to the left or to the right as soon as it left the starting base. Further, the data showed no evidence of on-line correction to the direction dimension of the task for the translated trials. Taken together, the results of the two experiments strongly suggest that, for fast video-aiming movements, the information concerning ones movement that is available in peripheral vision is used off-line.

Gaze influences finger movement-related and visual-related activation across the human brain.

The brain uses gaze orientation to organize myriad spatial tasks including hand movements. However, the neural correlates of gaze signals and their interaction with brain systems for arm movement control remain unresolved. Many studies have shown that gaze orientation modifies neuronal spike discharge in monkeys and activation in humans related to reaching and finger movements in parietal and frontal areas. To continue earlier studies that addressed interaction of horizontal gaze and hand movements in humans (Baker et al. 1999), we assessed how horizontal and vertical gaze deviations modified finger-related activation, hypothesizing that areas throughout the brain would exhibit movement-related activation that depended on gaze angle. The results indicated finger movement-related activation related to combinations of horizontal, vertical, and diagonal gaze deviations. We extended our prior findings to observation of these gaze-dependent effects in visual cortex, parietal cortex, motor, supplementary motor area, putamen, and cerebellum. Most significantly, we found a modulation bias for increased activation toward rightward, upper-right and vertically upward gaze deviations. Our results indicate that gaze modulation of finger movement-related regions in the human brain is spatially organized and could subserve sensorimotor transformations.

Investigating brain connectivity using mixed effects vector autoregressive models.

We propose a mixed-effects vector auto-regressive (ME-VAR) model for studying brain effective connectivity. One common approach to investigating inter-regional associations in brain activity is the multivariate auto-regressive (VAR) model. The standard VAR model unrealistically assumes the connectivity structure to be identical across all participants in a study and therefore, could yield misleading results. The ME-VAR model overcomes this limitation by incorporating a participant-specific connectivity structure. In addition, the ME-VAR models can capture connectivity differences across experimental conditions and patient groups. The ME-VAR model directly decomposes the connectivity matrices into (i.) the condition-specific connectivity matrix, which is shared by all participants in the study (fixed effect) and (ii.) a participant-specific component (random effect) which accounts for between-subject variation in connectivity. An advantage of our approach is that it permits the use of both theoretical results on mixed effects models and existing statistical software when fitting the model. Another advantage of the proposed approach is that it provides improved estimates of the within-subject coefficients (the random effects) by pooling information across subjects in a single-stage rather than the usual two-stage approach. We illustrate the ME-VAR model on a functional MRI data set obtained to investigate brain connectivity in the prefrontal, pre-motor and parietal cortices while humans performed a motor-related, decision-making and action selection task.

Attention modulates generalization of visuomotor adaptation

Generalization represents the ability to transfer what has been learned in one context to another context beyond limited experience. Because acquired motor representations often have to be reinstated in a different or novel environment, generalization is a crucial part of visuomotor learning. In daily life, training for new motor skills often occurs in a complex environment, in which dividing attentional resources for multiple stimuli is required. However, it is unknown how dividing attention during learning affects the generalization of visuomotor learning. We examined how divided attention during training modulates the generalization of visuomotor rotational adaptation. Participants were trained to adapt to one direction with or without dividing attention to a simultaneously presented visual detection task. Then, they had to generalize rotational adaptation to other untrained directions. We show that visuomotor training with divided attention multiplicatively reduces the gain and sharpens the tuning of the generalization function. We suggest that limiting attention narrowly restricts an internal model, reducing the range and magnitude of transfer. This result suggests that attention modulates a selective subpopulation of neurons in motor areas, those with directional tuning values in or near the training direction.

Brain representations for acquiring and recalling visual-motor adaptations.

Humans readily learn and remember new motor skills, a process that likely underlies adaptation to changing environments. During adaptation, the brain develops new sensory-motor relationships, and if consolidation occurs, a memory of the adaptation can be retained for extended periods. Considerable evidence exists that multiple brain circuits participate in acquiring new sensory-motor memories, though the networks engaged in recalling these and whether the same brain circuits participate in their formation and recall have less clarity. To address these issues, we assessed brain activation with functional MRI while young healthy adults learned and recalled new sensory-motor skills by adapting to world-view rotations of visual feedback that guided hand movements. We found cerebellar activation related to adaptation rate, likely reflecting changes related to overall adjustments to the visual rotation. A set of parietal and frontal regions, including inferior and superior parietal lobules, premotor area, supplementary motor area and primary somatosensory cortex, exhibited non-linear learning-related activation that peaked in the middle of the adaptation phase. Activation in some of these areas, including the inferior parietal lobule, intra-parietal sulcus and somatosensory cortex, likely reflected actual learning, since the activation correlated with learning after-effects. Lastly, we identified several structures having recall-related activation, including the anterior cingulate and the posterior putamen, since the activation correlated with recall efficacy. These findings demonstrate dynamic aspects of brain activation patterns related to formation and recall of a sensory-motor skill, such that non-overlapping brain regions participate in distinctive behavioral events.

Allocation of attention for dissociated visual and motor goals

In daily life, selecting an object visually is closely intertwined with processing that object as a potential goal for action. Since visual and motor goals are typically identical, it remains unknown whether attention is primarily allocated to a visual target, a motor goal, or both. Here, we dissociated visual and motor goals using a visuomotor adaptation paradigm, in which participants reached toward a visual target using a computer mouse or a stylus pen, while the direction of the cursor was rotated 45° counter-clockwise from the direction of the hand movement. Thus, as visuomotor adaptation was accomplished, the visual target was dissociated from the movement goal. Then, we measured the locus of attention using an attention-demanding rapid serial visual presentation (RSVP) task, in which participants detected a pre-defined visual stimulus among the successive visual stimuli presented on either the visual target, the motor goal, or a neutral control location. We demonstrated that before visuomotor adaptation, participants performed better when the RSVP stream was presented at the visual target than at other locations. However, once visual and motor goals were dissociated following visuomotor adaptation, performance at the visual and motor goals was equated and better than performance at the control location. Therefore, we concluded that attentional resources are allocated both to visual target and motor goals during goal-directed reaching movements.