Vibrissal motor cortex in the rat: connections with the barrel field

Abstract

The flow of information in the sensorimotor cortex may determine how somatic information modulates motor cortex neuronal activity during voluntary movement. Electrophysiological recordings and neuroanatomical tracing techniques were used to study the connections between the primary somatosensory cortex (SI) and the vibrissal representation of the primary motor cortex (MI) in rodents. Intracortical microstimulation (ICMS) was applied to the vibrissal region of the motor cortex to identify a site from which stimulation evoked movements of the vibrissae. Movements of only a single whisker were evoked by applying low-intensity stimulating current to particular locations within MI. A single injection of either horseradish peroxidase (HRP) or biocytin was made at the stimulus site in each animal, to retrogradely label cells in the somatosensory cortex. Receptive field (RF) responses were recorded from neurons in the barrel cortex to identify the sensory cortex representation of the same whisker that responded to ICMS. The site at which neurons responded predominately to manual stimulation of this particular vibrissa was marked by a small electrolytic lesion. The projection from the somatosensory cortex to the identified whisker representation in the motor cortex was determined by mapping the location of labeled neurons in tissue sections processed for either HRP or biocytin. The relationship of the labeled cells in SI to the barrel structures was determined from adjacent sections that were stained for cytochrome oxidase. In all cases, the barrel column associated with the relevant whisker contained labeled cells. Surrounding barrels also contained labeled cells, although fewer in number. Very few labeled cells were found in non-contiguous barrels. These results show that the SI to MI projection is somatotopically arranged, such that the sensory cortex representation of a whisker is morphologically connected to the motor cortex representation of the same whisker. Thus, sensory information is relayed to MI from the relevant whisker region in SI. Adjacent whisker regions also appear to relay somatic input, but presumably to a lesser degree. A second group of animals received single small injections of the anterograde tracer, Phaseolus vulgaris leucoagglutinin, to an electrophysiologically identified whisker representation in the sensory cortex. A single narrow column of labeled fibers was found in the motor cortex following such injections. Thus, the sensory cortex appears to relay somatic information from the vibrissae to restricted regions of the motor cortex in a somatotopically organized manner. Furthermore, the stimulus-evoked whisker movements suggest that certain features of the output map of the motor cortex are discretely organized. These input/output relationships suggest that complex information processing within the vibrissal sensorimotor cortex is highly organized.