Premotor Cortex - Re-emergence of The Premotor Cortex

Re-emergence of The Premotor Cortex

The hypothesis of a separate premotor cortex re-emerged and gained ground in the 1980s. Several key lines of research helped to establish the premotor cortex by showing that it had properties distinct from those of the adjacent primary motor cortex.

Roland and colleagues studied the dorsal premotor cortex and the supplementary motor area in humans while blood flow in the brain was monitored in a positron emission scanner. When people made complex sensory-guided movements such as following verbal instructions, more blood flow was measured in the dorsal premotor cortex. When people made internally paced sequences of movements, more blood flow was measured in the supplementary motor area. When people made simple movements that required little planning, such as palpating an object with the hand, the blood flow was more limited to the primary motor cortex. By implication, the primary motor cortex was more involved in execution of simple movement, the premotor cortex was more involved in sensory guided movement, and the supplementary motor area was more involved in internally generated movements.

Wise and his colleagues studied the dorsal premotor cortex of monkeys. The monkeys were trained to perform a delayed response task, making a movement in response to a sensory instruction cue. During the task, neurons in the dorsal premotor cortex became active in response to the sensory cue and often remained active during the few seconds of delay or preparation time before the monkey performed the instructed movement. Neurons in the primary motor cortex showed much less activity during the preparation period and were more likely to be active only during the movement itself. By implication, the dorsal premotor cortex was more involved in planning or preparing for movement and the primary motor cortex more involved in executing movement.

Rizzolatti and colleagues divided the premotor cortex into four parts or fields based on cytoarchitectonics, two dorsal fields and two ventral fields. They then studied the properties of the ventral premotor fields, establishing tactile, visual, and motor properties of a complex nature (summarized in greater detail below in Divisions of the premotor cortex).

At least three representations of the hand were reported in the motor cortex, one in the primary motor cortex, one in the ventral premotor cortex, and one in the dorsal premotor cortex. By implication, at least three different cortical fields may exist, each one performing its own special function in relation to the fingers and wrist.

For these and other reasons, a consensus has now emerged that the lateral motor cortex does not consist of a single, simple map of the body, but instead contains multiple subregions including the primary motor cortex and several premotor fields. These premotor fields have diverse properties. Some project to the spinal cord and may play a direct role in movement control, whereas others do not. Whether these cortical areas are arranged in a hierarchy or share some other more complex relationship is still debated.

Graziano and colleagues suggested an alternative principle of organization for the primary motor cortex and the caudal part of the premotor cortex, all regions that project directly to the spinal cord and that were included in the Penfield and Woolsey definition of M1. In this alternative proposal, the motor cortex is organized as a map of the natural behavioral repertoire. The complicated, multifaceted nature of the behavioral repertoire results in a complicated, heterogeneous map in cortex, in which different parts of the movement repertoire are emphasized in different cortical subregions. More complex movements such as reaching or climbing require more coordination among body parts, the processing of more complex control variables, the monitoring of objects in the space near the body, and planning several seconds into the future. Other parts of the movement repertoire, such as manipulating an object with the fingers once the object has been acquired, or manipulating an object in the mouth, involve less planning, less computation of spatial trajectory, and more control of individual joint rotations and muscle forces. In this view the more complex movements, especially multi-segmental movements, come to be emphasized in the more anterior part of the motor map because that cortex emphasizes the musculature of the back and neck which serves as the coordinating link between body parts. In contrast the simpler parts of the movement repertoire that tend to focus more on the distal musculature are emphasized in the more posterior cortex. In this alternative view, though movements of lesser complexity are emphasized in the primary motor cortex and movements of greater complexity are emphasized in the caudal premotor cortex, this difference does not necessarily imply a control hierarchy. Instead the regions differ from each other, and contain subregions with differing properties, because the natural movement repertoire itself is heterogeneous.