Neural coding of action in three dimensions: Task- and time-invariant reference frames for visuospatial and motor-related activity in parietal area V6A.

Goal-directed movements involve a series of neural computations that compare the sensory representations of goal location and effector position, and transform these into motor commands. Neurons in posterior parietal cortex (PPC) control several effectors (e.g., eye, hand, foot) and encode goal location in a variety of spatial coordinate systems, including those anchored to gaze direction, and to the positions of the head, shoulder, or hand. However, there is little evidence on whether reference frames depend also on the effector and/or type of motor response. We addressed this issue in macaque PPC area V6A, where previous reports using a fixate-to-reach in depth task, from different starting arm positions, indicated that most units use mixed body/hand-centered coordinates. Here, we applied singular value decomposition and gradient analyses to characterize the reference frames in V6A while the animals, instead of arm reaching, performed a nonspatial motor response (hand lift). We found that most neurons used mixed body/hand coordinates, instead of "pure" body-, or hand-centered coordinates. During the task progress the effect of hand position on activity became stronger compared to target location. Activity consistent with body-centered coding was present only in a subset of neurons active early in the task. Applying the same analyses to a population of V6A neurons recorded during the fixate-to-reach task yielded similar results. These findings suggest that V6A neurons use consistent reference frames between spatial and nonspatial motor responses, a functional property that may allow the integration of spatial awareness and movement control.

Mixed Body/Hand Reference Frame for Reaching in 3D Space in Macaque Parietal Area PEc.

The neural correlates of coordinate transformations from vision to action are expressed in the activity of posterior parietal cortex (PPC). It has been demonstrated that among the medial-most areas of the PPC, reaching targets are represented mainly in hand-centered coordinates in area PE, and in eye-centered, body-centered, and mixed body/hand-centered coordinates in area V6A. Here, we assessed whether neurons of area PEc, located between V6A and PE in the medial PPC, encode targets in body-centered, hand-centered, or mixed frame of reference during planning and execution of reaching. We studied 104 PEc cells in 3 Macaca fascicularis. The animals performed a reaching task toward foveated targets located at different depths and directions in darkness, starting with the hand from 2 positions located at different depths, one next to the trunk and the other far from it. We show that most PEc neurons encoded targets in a mixed body/hand-centered frame of reference. Although the effect of hand position was often rather strong, it was not as strong as reported previously in area PE. Our results suggest that area PEc represents an intermediate node in the gradual transformation from vision to action that takes place in the reaching network of the dorsomedial PPC.

Overlapping representations for reach depth and direction in caudal superior parietal lobule of macaques.

Reaching movements in the real world have typically a direction and a depth component. Despite numerous behavioral studies, there is no consensus on whether reach coordinates are processed in separate or common visuomotor channels. Furthermore, the neural substrates of reach depth in parietal cortex have been ignored in most neurophysiological studies. In the medial posterior parietal area V6A, we recently demonstrated the strong presence of depth signals and the extensive convergence of depth and direction information on single neurons during all phases of a fixate-to-reach task in 3-dimensional (3D) space. Using the same task, in the present work we examined the processing of direction and depth information in area PEc of the caudal superior parietal lobule (SPL) in three Macaca fascicularis monkeys. Across the task, depth and direction had a similar, high incidence of modulatory effect. The effect of direction was stronger than depth during the initial fixation period. As the task progressed toward arm movement execution, depth tuning became more prominent than directional tuning and the number of cells modulated by both depth and direction increased significantly. Neurons tuned by depth showed a small bias for far peripersonal space. Cells with directional modulations were more frequently tuned toward contralateral spatial locations, but ipsilateral space was also represented. These findings, combined with results from neighboring areas V6A and PE, support a rostral-to-caudal gradient of overlapping representations for reach depth and direction in SPL. These findings also support a progressive change from visuospatial (vergence angle) to somatomotor representations of 3D space in SPL.