Spatiotemporal dynamics of cortical sensorimotor integration in behaving mice.

Long-range neuronal circuits underlying the interaction between sensory and motor cortex.

The primary somatosensory barrel cortex processes tactile vibrissae information, allowing rodents to actively perceive spatial and textural features of their immediate surroundings. Each whisker on the snout is individually represented in the neocortex by an anatomically identifiable 'barrel' specified by the segregated termination zones of thalamocortical axons of the ventroposterior medial nucleus, which provide the primary sensory input to the neocortex. The sensory information is subsequently processed within local synaptically connected neocortical microcircuits, which have begun to be investigated in quantitative detail. In addition to these local synaptic microcircuits, the excitatory pyramidal neurons of the barrel cortex send and receive long-range glutamatergic axonal projections to and from a wide variety of specific brain regions. Much less is known about these long-range connections and their contribution to sensory processing. Here, we review current knowledge of the long-range axonal input and output of the mouse primary somatosensory barrel cortex. Prominent reciprocal projections are found between primary somatosensory cortex and secondary somatosensory cortex, motor cortex, perirhinal cortex and thalamus. Primary somatosensory barrel cortex also projects strongly to striatum, thalamic reticular nucleus, zona incerta, anterior pretectal nucleus, superior colliculus, pons, red nucleus and spinal trigeminal brain stem nuclei. These long-range connections of the barrel cortex with other specific cortical and subcortical brain regions are likely to play a crucial role in sensorimotor integration, sensory perception and associative learning.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1460-9568.2010.07264.x

Long-range connectivity of mouse primary somatosensory barrel cortex.

Recent techniques for tracing pathways in the central nervous system of developing and adult mammals.

Pharmacological inactivation in the analysis of the central control of movement.

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.

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

Direct corticocortical afferents to the primary motor cortex (MI) originate in area 2 and area 3a of the primary somatosensory cortex (SI). The functional and morphological characteristics of the two pathways indicate that they relay different sensory signals to MI. The role of area 2 in relaying peripheral information to the cat MI was studied using electrophysiological techniques. Neurons that responded to stimulation of peripheral receptive fields on the contralateral forepaw were identified in MI by extracellular recordings. In area 2 of SI, neurons with the same receptive field modality and location as those in MI were also identified. Field potentials to electrical stimulation of the peripheral receptive field were recorded at the somatotopically matched sites in both MI and SI. Neuronal activity at the recording site in area 2 was blocked by injection of lidocaine, a local anesthetic. Changes in MI and area 2 responses were monitored before and after inactivation of area 2. Neuronal activity near t...

The Effects of Localized Inactivation of Somatosensory Cortex, Area 2, on the Cat Motor Cortex

In cats and primates, area 3a of the somatosensory cortex is the primary recipient of proprioceptive input (Phillips et al., 1971). Neurons in area 3a project to area 2 (Pons and Kaas, 1986; Porter, 1991), where somatic input relayed from the cortex and the thalamus may be integrated (Iwamura and Tanaka, 1978a,c). The goal of the present study was to determine the effects of area 3a input on neuronal activity in area 2 of cats. Extracellular recording techniques were used to identify neurons in area 2 that responded to deep stimulation of the contralateral forepaw. Neurons in area 3a that responded to the same receptive field modality and location as those in area 2 were also isolated. Single-unit or multiunit responses and evoked potentials to electrical stimulation of the shared peripheral receptive field and spontaneous activity were recorded from areas 2 and 3a. Lidocaine, a local anesthetic, was injected at the area 3a recording site to block neuronal activity. Spontaneous activity and receptive fields were abolished and evoked potentials were considerably diminished at the injection site, immediately after lidocaine was administered. Changes in unit responses, spontaneous activity, and evoked potentials in area 2 were monitored following inactivation of the somatotopically "matched" site in area 3a. Unit activity was recorded at 15 matched sites. In area 2, changes in unit responses to the peripheral stimulation and/or in spontaneous activity were observed at most of the recording sites following inactivation of area 3a. Spontaneous activity rates changed at 63% of the sites (mean change = 85%). Unit responses to the peripheral stimulation changed at 57% of the recording sites (mean change = 47%). The remaining sites in area 2 did not show lidocaine-induced changes. These sites may not have been connected with the matched sites in area 3a. Spontaneous activity and unit responses were not always similarly altered at a given site; sometimes one increased while the other decreased. Decreases in unit responses and spontaneous activity following inactivation of area 3a input were the predominant effects, indicating that area 3a has a facilitatory effect on neuronal activity observed in some regions of area 2. However, increases in neuronal activity at some sites indicated that the effects of area 3a input on area 2 are nonuniform. Evoked potentials were recorded at 19 matched sites, before and after injection of lidocaine. Evoked potentials also changed at some area 2 recording sites following area 3a inactivation.(ABSTRACT TRUNCATED AT 400 WORDS)

Ruth Izraeli

Papers

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

The Effects of Localized Inactivation of Somatosensory Cortex, Area 2, on the Cat Motor Cortex

The Effects of Localized Inactivation of Somatosensory Cortex, Area 3a, on Area 2 in Cats