Cortical Mechanisms for Reaches Versus Saccades: Progression of Effector-Specificity Through Target Memory to Movement Planning and Execution

Abstract

Effector-specific cortical mechanisms can be difficult to establish using fMRI, in part because low time resolution might temporally conflate different signals related to target representation, motor planning, and motor execution. Here, we used an event-related fMRI protocol and a cue-separation paradigm to temporally separate these three major sensorimotor stages for saccades vs. reaches. In each trial, subjects (N=12) 1) briefly viewed a target 4-7{degrees} left or right of midline fixation on a touchscreen, followed by an 8 second delay (effector-independent target memory phase), 2) were instructed by an auditory cue to perform a reach or a saccade, followed by a second delay of 8 seconds (effector-specific planning phase), and finally 3) were prompted to move by reaching-to-touch or performing a saccade towards the remembered target (effector-specific execution phase). Our analysis of saccade and reach activation (vs. a non-spatial control task) revealed modest effector-agnostic target memory activity (left AG, bilateral mIPS) followed by independent effector parietofrontal sites and time courses during the motor components of the task, specifically: more medial (pIPS, mIPS, M1, and PMd) activity during both reach planning and execution, and more lateral (mIPS, AG, and FEF) activity only during saccade execution. These motor activations were bilateral, with a left (contralateral) preference for reach. A conjunction analysis revealed that left mIPS and right AG, PCu, SPOC, FEF/PMv and LOTC showed activation for both saccades and reaches. Overall, effector-preference contrasts (reach vs. saccade) revealed significantly more parietofrontal activation for reaches than saccades during both planning and execution, with the exception of FEF. Cross-correlation of reach, saccade, and reach-saccade activation through time revealed correlated activation both within and across effectors in each hemisphere, but with a tendency toward higher correlations in the right hemisphere, especially between the eye and hand. These results demonstrate substantially independent but temporally correlated cortical networks for human eye, hand, and eye-hand control, that follow explicit spatiotemporal rules for effector-specific timing, medial-lateral distribution, and hemispheric lateralization.