Maintained Representations of the Ipsilateral and Contralateral Limbs during Bimanual Control in Primary Motor Cortex.

Primary motor cortex (M1) almost exclusively controls the contralateral side of the body. However, M1 activity is also modulated during ipsilateral body movements. Previous work has shown that M1 activity related to the ipsilateral arm is independent of the M1 activity related to the contralateral arm. How do these patterns of activity interact when both arms move simultaneously? We explored this problem by training 2 monkeys (male, Macaca mulatta) in a postural perturbation task while recording from M1. Loads were applied to one arm at a time (unimanual) or both arms simultaneously (bimanual). We found 83% of neurons (n = 236) were responsive to both the unimanual and bimanual loads. We also observed a small reduction in activity magnitude during the bimanual loads for both limbs (25%). Across the unimanual and bimanual loads, neurons largely maintained their preferred load directions. However, there was a larger change in the preferred loads for the ipsilateral limb (∼25%) than the contralateral limb (∼9%). Lastly, we identified the contralateral and ipsilateral subspaces during the unimanual loads and found they captured a significant amount of the variance during the bimanual loads. However, the subspace captured more of the bimanual variance related to the contralateral limb (97%) than the ipsilateral limb (66%). Our results highlight that, even during bimanual motor actions, M1 largely retains its representations of the contralateral and ipsilateral limbs.SIGNIFICANCE STATEMENT Previous work has shown that primary motor cortex (M1) represents information related to the contralateral limb, its downstream target, but also reflects information related to the ipsilateral limb. Can M1 still represent both sources of information when performing simultaneous movements of the limbs? Here we record from M1 during a postural perturbation task. We show that activity related to the contralateral limb is maintained between unimanual and bimanual motor actions, whereas the activity related to the ipsilateral limb undergoes a small change between unimanual and bimanual motor actions. Our results indicate that two independent representations can be maintained and expressed simultaneously in M1.

Independent representations of ipsilateral and contralateral limbs in primary motor cortex.

Several lines of research demonstrate that primary motor cortex (M1) is principally involved in controlling the contralateral side of the body. However, M1 activity has been correlated with both contralateral and ipsilateral limb movements. Why does ipsilaterally-related activity not cause contralateral motor output? To address this question, we trained monkeys to counter mechanical loads applied to their right and left limbs. We found >50% of M1 neurons had load-related activity for both limbs. Contralateral loads evoked changes in activity ~10ms sooner than ipsilateral loads. We also found corresponding population activities were distinct, with contralateral activity residing in a subspace that was orthogonal to the ipsilateral activity. Thus, neural responses for the contralateral limb can be extracted without interference from the activity for the ipsilateral limb, and vice versa. Our results show that M1 activity unrelated to downstream motor targets can be segregated from activity related to the downstream motor output.

Primary motor cortex neurons classified in a postural task predict muscle activation patterns in a reaching task.

Primary motor cortex (M1) activity correlates with many motor variables, making it difficult to demonstrate how it participates in motor control. We developed a two-stage process to separate the process of classifying the motor field of M1 neurons from the process of predicting the spatiotemporal patterns of its motor field during reaching. We tested our approach with a neural network model that controlled a two-joint arm to show the statistical relationship between network connectivity and neural activity across different motor tasks. In rhesus monkeys, M1 neurons classified by this method showed preferred reaching directions similar to their associated muscle groups. Importantly, the neural population signals predicted the spatiotemporal dynamics of their associated muscle groups, although a subgroup of atypical neurons reversed their directional preference, suggesting a selective role in antagonist control. These results highlight that M1 provides important details on the spatiotemporal patterns of muscle activity during motor skills such as reaching.

Designing a thalamic somatosensory neural prosthesis: consistency and persistence of percepts evoked by electrical stimulation.

Intuitive somatosensory feedback is required for fine motor control. Here we explored whether thalamic electrical stimulation could provide the necessary durations and consistency of percepts for a human somatosensory neural prosthetic. Continuous and cycling high-frequency (185 Hz, 0.21 ms pulse duration charge balanced square wave) electrical pulses with the cycling patterns varying between 7% and 67% of duty cycle were applied in five patients with chronically implanted deep brain stimulators. Stimulation produced similar percepts to those elicited immediately after surgery. While consecutive continuous stimuli produced decreasing durations of sensation, the amplitude and type of percept did not change. Cycling stimulation with shorter duty cycles produced more persisting percepts. These features suggest that the thalamus could provide a site for stable and enduring sensations necessary for a long term somatosensory neural prosthesis.