On the functional organization and operational principles of the motor cortex.

Recent studies on the functional organization and operational principles of the motor cortex (MCx), taken together, strongly support the notion that the MCx controls the muscle synergies subserving movements in an integrated manner. For example, during pointing the shoulder, elbow and wrist muscles appear to be controlled as a coupled functional system, rather than singly and separately. The recurrent pattern of intrinsic synaptic connections between motor cortical points is likely part of the explanation for this operational principle. So too is the reduplicated, non-contiguous and intermingled representation of muscles in the MCx. A key question addressed in this article is whether the selection of movement related muscle synergies is a dynamic process involving the moment to moment functional linking of a variety of motor cortical points, or rather the selection of fixed patterns embedded in the MCx circuitry. It will be suggested that both operational principles are probably involved. We also discuss the neural mechanisms by which cortical points may be dynamically linked to synthesize movement related muscle synergies. Separate corticospinal outputs sum linearly and lead to a blending of the movements evoked by activation of each point on its own. This operational principle may simplify the synthesis of motor commands. We will discuss two possible mechanisms that may explain linear summation of outputs. We have observed that the final posture of the arm when pointing to a given spatial location is relatively independent of its starting posture. From this observation and the recurrent nature of the MCx intrinsic connectivity we hypothesize that the basic mode of operation of the MCx is to associate spatial location to final arm posture. We explain how the recurrent network connectivity operates to generate the muscle activation patterns (synergies) required to move the arm and hold it in its final position.