The genitals and gluteal skin are represented lateral to the foot in anterior parietal somatosensory cortex of macaques

Detailed electrophysiological maps of the representations of trunk and adjacent body parts in area 3b and area 1 of somatosensory cortex were obtained in three macaque monkeys ( Macaca mulatta and Macaca radiata ) of either sex. A total of 211 microelectrode penetrations 250-300 &#119 m apart resulted in 1,190 recording sites. During penetrations deep into the posterior bank of the central sulcus, recordings were made every 300 &#119 m to depths of 6-7 mm until sites unresponsive to somatic stimuli were reached. Cortex was later cut parasagittally and sections were stained for cytochrome oxidase (CO) or Nissl substance. Contrary to expectations from earlier reports, the genitalia were represented lateral to the representations of the foot in cortex along the area 3b/1 border. The gluteal skin including the gluteal pads and the base of the tail were also represented in this section of cortex. Only a small region of cortex was devoted to the genitalia, and neurons in this cortex had receptive fields that were large and typically included skin of the inner thigh and belly. The lower, middle and upper trunk were represented more laterally, followed by the neck, upper head and arm. The receptive fields on the trunk were roughly the same size as those for the middle and lower trunk and slightly smaller on the upper trunk.     

The Corticocuneate Pathway in the Cat: Relations among Terminal Distribution Patterns,Cytoarchitecture, and Single Neuron Functional Properties

A combined anatomical and physiological strategy was used to investigate the organization of the corticocuneate pathway in the cat. The distribution of the corticocuneate projection was mapped by means of the anterograde horseradish peroxidase (HRP) labeling technique and correlated with the nuclear cytoarchitecture in Nissl and Golgi material, the distribution of retrogradely labeled relay cells after HRP injections in the ventrobasal complex of the thalamus, and the topographic organization derived from single-and multiunit recordings in the decerebrate, unanesthetized cat. This approach provided details about the arrangement of the corticocuneate pathway that were not available from previous studies with anterograde degeneration methods.

On the basis of cytoarchitectonic and connectional features, a number of subdivisions are identified in the cuneate nucleus, each of which is associated with characteristic functional properties. In agreement with previous studies, it is found that a large portion of the cuneate nucleus, the middle dorsal part (MCd), is exclusively devoted to the representation of cutaneous receptive fields on the digits. This “core” region contains more thalamic projecting neurons than any other subdivision of the cuneate nucleus. A topographic arrangement also exists in the subdivisions of the rostral cuneate and of the nuclear region ventral to MCd, although in these, receptive fields are larger and predominantly, but not exclusively, related to deep receptors and involve the arm, shoulder, and trunk.

Observations on corticocuneate projections were based on injections, mainly focused on functional subdivisions of the primary somatosensory cortex (SI) as described by McKenna et al (1981). Although cortical projections are mainly to cuneate regions other than its core, a significant proportion of fibers from the region of SI where the digits are represented (particularly area 3b) do project to the MCd region of the cuneate nucleus. Similarly, nuclear areas associated with receptive fields on the arm and trunk are labeled after injection in SI arm and trunk regions, respectively. Thus, a close topographic relationship appears to exist between the somatosensory cortex and cuneate regions related to the same body representation, although nuclear regions in which receptive fields on the neck area are represented receive very sparse or no detectable cortical projections even when the injection of the tracer involves the entire sensorimotor cortex. The topographic arrangement of SI projections upon the cuneate nucleus suggests that a similar pattern exists in both structures with regard to the relative representations of distal versus proximal and deep versus cutaneous receptive fields (e.g., “core” vs. “shell” organization), and that cuneate regions preferentially related to either of these classes of receptive fields receive direct connections from the corresponding regions in SI.

A comparison of the results from cats with tracer injections in areas 4 and 3b reveals that the projections from the former is denser than that arising from the latter and that their territories of termination largely overlap in the ventral portions of the cuneate nucleus. However, cortical projections to MCd may be derived from the somatosensory cortex with no contribution from area 4. The demonstration of the relative selectivity of cortical projections from different cytoarchitectonic and functional cortical areas to cuneate regions identified here provides a structural basis for the elucidation of the physiological and behavioral observations, particularly on cortical modulation of somatosensory transmission during movements.     

Differences in orientation and receptive field position between supra- and infragranular cells of cat striate cortex and their possible functional implications

On the postlateral gyrus of the cat striate cortex the cells' preferred orientation and the location of their receptive fields was measured as a function of cortical depth in penetrations as parallel as possible to the radiating fibres. In most penetrations the majority of infragranular cells showed orientation preferences 45 degrees-90 degrees different from the preferred orientations of supragranular cells. In addition, aggregate receptive fields from the same eye of supra- and infragranular cells were spatially shifted against each other. Using different columnar models these results are discussed in terms of spatial contrast enhancement for two parallel mechanisms in upper and lower layers, determined for pattern discrimination and movement detection.     

Sensory representation abnormalities that parallel focal hand dystonia in a primate model

In our hypothesis of focal dystonia, attended repetitive behaviors generate aberrant sensory representations. Those aberrant representations interfere with motor control. Abnormal motor control strengthens sensory abnormalities. The positive feedback loop reinforces the dystonic condition. Previous studies of primates with focal hand dystonia have demonstrated multi-digit or hairy-glabrous responses at single sites in area 3b, receptive fields that average ten times larger than normal, and high receptive field overlap as a function of horizontal distance. In this study, we strengthen and elaborate these findings. One animal was implanted with an array of microelectrodes that spanned the border between the face and digits. After the animal developed hand dystonia, responses in the initial hand representation increasingly responded to low threshold stimulation of the face in a columnar substitution. The hand-face border that is normally sharp became patchy and smeared over 1 mm of cortex within 6 weeks. Two more trained animals developed a focal hand dystonia variable in severity across the hand. Receptive field size, presence of multi-digit or hairy-glabrous receptive fields, and columnar overlap covaried with the animal's ability to use specific digits. A fourth animal performed the same behaviors without developing dystonia. Many of its physiological measures were similar to the dystonic animals, but receptive field overlap functions were minimally abnormal, and no sites shared response properties that are normally segregated such as hairy-glabrous combined fields, or multi-digit fields. Thalamic mapping demonstrated proportionate levels of abnormality in thalamic representations as were found in cortical representations.     

Sensory representation abnormalities that parallel focal hand dystonia in a primate model

In our hypothesis of focal dystonia, attended repetitive behaviors generate aberrant sensory representations. Those aberrant representations interfere with motor control. Abnormal motor control strengthens sensory abnormalities. The positive feedback loop reinforces the dystonic condition. Previous studies of primates with focal hand dystonia have demonstrated multi-digit or hairy-glabrous responses at single sites in area 3b, receptive fields that average ten times larger than normal, and high receptive field overlap as a function of horizontal distance. In this study, we strengthen and elaborate these findings. One animal was implanted with an array of microelectrodes that spanned the border between the face and digits. After the animal developed hand dystonia, responses in the initial hand representation increasingly responded to low threshold stimulation of the face in a columnar substitution. The hand-face border that is normally sharp became patchy and smeared over 1 mm of cortex within 6 weeks. Two more trained animals developed a focal hand dystonia variable in severity across the hand. Receptive field size, presence of multi-digit or hairy-glabrous receptive fields, and columnar overlap covaried with the animal's ability to use specific digits. A fourth animal performed the same behaviors without developing dystonia. Many of its physiological measures were similar to the dystonic animals, but receptive field overlap functions were minimally abnormal, and no sites shared response properties that are normally segregated such as hairy-glabrous combined fields, or multi-digit fields. Thalamic mapping demonstrated proportionate levels of abnormality in thalamic representations as were found in cortical representations.     

Ablations of areas 3b (SI proper) and 3a of somatosensory cortex in marmosets deactivate the second and parietal ventral somatosensory areas

Partial ablations of specific parts of cortical areas 3b (SI proper) and 3a in marmosets were found to render somatotopically equivalent parts of two other cortical somatosensory fields, the second somatosensory area (SII) and the parietal ventral area (PV), unresponsive to peripheral stimulation. Microelectrode recordings in anesthetized marmosets first established the responsiveness and locations of the representations of body parts, including the hand in areas 3a and 3b, SII, and in some cases PV. The hand representations in areas 3a and 3b were then removed by aspiration. Immediately afterwards, additional recordings established that regions of SII and PV that formerly represented the hand were no longer responsive to cutaneous stimulation of the hand (or any other skin surface). Other parts of these fields, representing parts of the body other than the hand, remained responsive to stimulation of the previously effective receptive fields. We conclude that SII and PV depend on inputs (either direct or indirect) from areas 3a and 3b for their activation.     

Experience-induced plasticity of cutaneous maps in the primary somatosensory cortex of adult monkeys and rats

In a first study, the representations of skin surfaces of the hand in the primary somatosensory cortex, area 3b, were reconstructed in owl monkeys and squirrel monkeys trained to pick up food pellets from small, shallow wells, a task which required skilled use of the digits. Training sessions included limited manual exercise over a total period of a few hours of practice. From an early clumsy performance in which many retrieval attempts were required for each successful pellet retrieval, the monkeys exhibited a gradual improvement. Typically, the animals used various combinations of digits before developing a successful retrieval strategy. As the behavior came to be stereotyped, monkeys consistently engaged surfaces of the distal phalanges of one or two digits in the palpation and capture of food pellets from the smallest wells. Microelectrode mapping of the hand surfaces revealed that the glabrous skin of the fingertips predominantly involved in the dexterity task was represented over topographically expanded cortical sectors. Furthermore, cutaneous receptive fields which covered the most frequently stimulated digital tip surfaces were less than half as large as were those representing the corresponding surfaces of control digits. In a second series of experiments, Long-Evans rats were assigned to environments promoting differential tactile experience (standard, enriched, and impoverished) for 80 to 115 days from the time of weaning. A fourth group of young adult rat experienced a severe restriction of forepaw exploratory movement for either 7 or 15 days. Cortical maps derived in the primary somatosensory cortex showed that environmental enrichment induced a substantial enlargement of the cutaneous forepaw representation, and improved its spatial resolution (smaller glabrous receptive fields). In contrast, tactile impoverishment resulted in a degradation of the forepaw representation that was characterized by larger cutaneous receptive fields and the emergence of non-cutaneous responses. Cortical maps derived in the hemispheres contralateral to the immobilized forelimb exhibited a severe decrease of about 50% in the overall areal extent of the cutaneous representation of the forepaw, which resulted from the invasion of topographically organized cortical zones of non-cutaneous responses, and numerous discontinuities in the representation of contiguous skin territories. The size and the spatial arrangement of the cutaneous receptive fields were not significantly modified by the immobilization of the contralateral forelimb. Similar results were obtained regardless of whether the forelimb restriction lasted 7 or 15 days. These two studies corroborate the view that representational constructs are permanently reshaped by novel experiences through dynamic competitive processes. These studies also support the notion that subject-environment interactions play a crucial role in the maintenance of basic organizational features of somatosensory representations.     

Topographic organization of baboon primary motor cortex: face, hand, forelimb, and shoulder representation

(1) The fine details of the motor organization of the forelimb, face, and tongue representation of the baboon (Papio h. anubis) primary motor cortex were studied in four adult animals, using intracortical microstimulation (ICMS). (2) A total of 293 electrode penetrations were made. ICMS was delivered to 10,052 sites, and of these, 6,186 sites were verified to have been located within the grey matter. Motor effects were evoked from 30% of these sites. (3) The baboon motor cortex is confined, in large part, to the cortical tissue lying along the anterior bank of the central sulcus. When the electrode penetrations were confined to the precentral gyrus, few sites were capable of evoking movement when stimulated by currents of 40 microA or less. (4) The details of the motor maps varied among the four animals; nonetheless, a general topographic organization existed, with the tongue musculature being represented most laterally, followed by a medial progression of the face, digits, wrist, forearm, and shoulder. Within the representation of a given body part, the muscles were organized as a mosaic, wherein the same muscle was multiply represented. (5) A zone of unresponsive cortex was observed to lie consistently between the face and forelimb representation in all four animals. Repeated electrode penetrations within the unresponsive zone failed to elicit muscle contractions even with stimulating currents as high as 80 microA. (6) Our results suggest that the baboon motor cortex is topographically organized; however, embedded within this overall pattern lies a fine-grained mosaic incorporating multiple representations of the same muscle.     

Topographic Organization of Baboon Primary Motor Cortex: Face,Hand, Forelimb,and Shoulder Representation

(1) The fine details of the motor organization of the forelimb, face, and tongue representation of the baboon (Papio h. anubis)primary motor cortex were studied in four adult animals, using intracortical microstimulation (ICMS). (2) A total of 293 electrode penetrations were made. ICMS was delivered to 10,052 sites, and of these, 6,186 sites were verified to have been located within the grey matter. Motor effects were evoked from 30% of these sites. (3)The baboon motor cortex is confined, in large part, to the cortical tissue lying along the anterior bank of the central sulcus. When the electrode penetrations were confined to the precentral gyrus, few sites were capable of evoking movement when stimulated by currents of 40 μA or less. (4)The details of the motor maps varied among the four animals; nonetheless, a general topographic organization existed, with the tongue musculature being represented most laterally, followed by a medial progression of the face, digits, wrist, forearm, and shoulder. Within the representation of a given body part, the muscles were organized as a mosaic, wherein the same muscle was multiply represented. (5) A zone of unresponsive cortex was observed to lie consistently between the face and forelimb representation in all four animals. Repeated electrode penetrations within the unresponsive zone failed to elicit muscle contractions even with stimulating currents as high as 80 μA. (6) Our results suggest that the baboon motor cortex is topographically organized; however, embedded within this overall pattern lies a fine-grained mosaic incorporating multiple representations of the same muscle.     

Receptive fields of auditory cortical neurons in the cat

Receptive fields of auditory cortical neurons were studied by electrical stimulation of nerve fibers in different parts of the cochlea in cats anesthetized with pentobarbital. The dimensions of the receptive fields were shown to depend on the topographic arrangement of the neuron in the auditory cortex. The more caudad the neuron on the cortical projection of the cochlea in the primary auditory cortex, the more extensive its receptive field. The receptive fields were narrowest in the basal turn of the cochlea and were symmetrical with respect to their center. It is suggested that the region of finest discrimination of acoustic stimuli in cats is located in the basal region of the cochlea, i.e., in that part of its receptor system which has the narrowest receptive field and is represented by significantly more (than the middle and apical regions of the cochlea) nerve cells in the primary auditory cortex [1].A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 5, pp. 467–473, September–October, 1981.     

Bilateral receptive field neurons and callosal connections in the somatosensory cortex

Earlier studies recording single neuronal activity with bilateral receptive fields in the primary somatosensory cortex of monkeys and cats agreed that the bilateral receptive fields were related exclusively to the body midline and that the ipsilateral information reaches the cortex via callosal connections since they are dense in the cortical region representing the midline structures of the body while practically absent in the regions representing the distal extremities. We recently found a substantial number of neurons with bilateral receptive fields on hand digits, shoulders-arms or legs-feet in the caudalmost part (areas 2 and 5) of the postcentral gyrus in awake Japanese monkeys (Macaca fuscata). I review these results, discuss the functional implications of this bilateral representation in the postcentral somatosensory cortex from a behavioural standpoint and give a new interpretation to the midline fusion theory.     

Somatotopic Organization of the Third Somatosensory Area (SIII) in Cats

Multiunit microelectrode recording techniques were used to study the location and organization of the third somatosensory area (SIII) in cats. Representations of all major contralateral body parts were found in a small region of cortex along the lateral wing of the ansate sulcus and between the lateral sulcus and the suprasylvian sulcus. The systematic map of the body surface included forepaw and face regions previously identified as parts of SIII. The forepaw representation was generally buried on the rostral bank of the lateral wing of the ansate sulcus. The representations of the face and mystacial vibrissae were largely exposed on the rostral suprasylvian gyrus, but part of the representation of the face was also buried in the lateral wing of the ansate sulcus. Representations of the trunk and hindlimb extended from the suprasylvian gyrus onto the medial bank of the suprasylvian sulcus. We had expected to find these latter body parts in more medial cortex just caudal to the representation of these parts in the first somatosensory area (SI). Instead, neurons in penetrations in cortex caudal to the SI trunk and hindlimb representations were unresponsive to tactile stimulation. The unexpected location of the hindlimb in SIII led us to determine whether the proposed parts of SIII had similar cortical and thalamic connections. Injected anatomical tracers revealed that the representations of both the forelimb and hindlimb were interconnected with SI and a region of the thalamus just dorsal to the ventroposterior nucleus. Similarities in patterns of connections of forelimb and hindlimb portions of SIII supported the conclusion that SHI as presented here is a functional unit of cortex. We conclude that SIII has a somatotopic organization that does not parallel that in SI, and that SIII is not entirely coextensive with either area 5 or area 5a of Hassler and Muhs-Clement (1964).     

Retinotopic organization of the posterior suprasylvian area of the feline cerebral cortex

Representation of the visual field was investigated in the feline posterior suprasylvian area (PSA) using electrophysiological mapping techniques. The PSA is one of the extrastriatal visual structures of the cerebral cortex. The PSA retinotopic organization pattern was also studied. Neuronal receptive fields (RF) were mainly located in the upper contralateral quadrant and just a small number in the lower contralateral quadrant of the visual field. Approximately 10% of RF were located in the upper ipsilateral quadrant. The central area of the visual field extending in a radius of 20–30° from the area centralis was mainly represented in the upper section of the PSA (areas 21a and 21b). The RP of neurons located more peripherally to the area centralis are found in the lower portion of the PSA (areas 20a and 20b); these occupy a correspondingly greater area. Experimental finding did not confirm any substantial differences in the retinotopic organization of areas 21a, 21b, 20a, and 20b comprising the PSA. Data obtained would tend to indicate that the PSA consists of two areas, 21a and 21b, which do not appear to be subdivided, with more densely distributed visual neurons in the former than in the latter.Institute of Experimental Biology of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 23, No. 3, pp. 290–296, May–June, 1991.     

The organization and connections of somatosensory cortex in the brush-tailed possum (Trichosurus vulpecula): evidence for multiple, topographically organized and interconnected representations in an Australian marsupial

Microelectrode mapping techniques were used to determine the organization of somatosensory cortex in the Australian brush-tailed possum (Trichosurus vulpecula). The results of electrophysiological mapping were combined with data on the cyto- and myeloarchitecture, and patterns of corticocortical connections, using sections cut tangential to the pial surface. We found evidence for three topographically organized representations of the body surface that were coextensive with architectonic subdivisions. A large, discontinuous cutaneous representation in anterior parietal cortex was termed the primary somatosensory area (SI). Lateral to SI we found evidence for two further areas, the second somatosensory area (SII) and the parietal ventral area (PV). While neurones in all of these areas were responsive to cutaneous stimulation, those of SI were non-habituating, whereas those in SII and PV often habituated to the stimuli. Moreover, neuronal receptive fields in SII and PV were, in general, larger than those in SI. Neurones in cortex adjacent to the rostral and caudal boundaries of SI, including cortex that interdigitated between the discontinuous SI head and body representations, required stimulation of deep receptors in the periphery to elicit responses. Within the region of cortex containing neurones responsive to stimulation of deep receptors, body parts were represented in a mediolateral progression. Injections of anatomical tracers placed in electrophysiologically identified locations in SI revealed ipsilateral connections with other parts of SI, as well as cortex rostral to, caudal to, and interdigitating between, SI. Injections in SI also resulted in labelling in PV, SII, motor cortex, posterior parietal cortex and perirhinal cortex. The patterns of contralateral projections reflected those of ipsilateral projections, although they were relatively less dense. The present findings support recent observations in other marsupials in which multiple representations of the body surface were described, and suggest that multiple interconnected sensory representations may be a common feature of cortical organization and function in marsupials.     

The organization of somatosensory cortex in the short-tailed opossum (Monodelphis domestica)

The organization of neocortex in the short-tailed opossum (Monodelphis domestica) was explored with multiunit microelectrode recordings from middle layers of cortex. Microelectrode maps were subsequently related to the chemoarchitecture of flattened cortical preparations, sectioned parallel to the cortical surface and processed for either cytochrome oxidase (CO) or NADPH-diaphorase (NADPHd) histochemistry. The recordings revealed the presence of at least two systematic representations of the contralateral body surface located in a continuous strip of cortex running from the rhinal sulcus to the medial wall. The primary somatosensory area (S1) was located medially while secondary somatosensory cortex (S2) formed a laterally located mirror image of S1. Auditory cortex was located in lateral cortex at the caudal border of S2, and some electrode penetrations in this area responded to both auditory and somatosensory stimulation. Auditory cortex was outlined by a dark oval visible in flattened brain sections. A large primary visual cortex (V1) was located at the caudal pole of cortex, and also consistently corresponded to a large chemoarchitecturally visible oval. Cortex just rostral and lateral to V1 responded to visual stimulation, while bimodal auditory/visual responses were obtained in an area between V1 and somatosensory cortex. The results are compared with brain organization in other marsupials and with placentals and the evolution of cortical areas in mammals is discussed.     

Responsiveness of reorganized primary somatosensory (SI) cortex after local inactivation of normal SI cortex in chronic spinal cats.

The cortical map of adult cats that sustained spinal cord transection at T12 when they were 2 weeks old is characterized by a clear duplication of the representation of the forelimb, rostral trunk, and neck. The novel representation is located in the cortical region that is, in nonoperated animals, normally devoted to the hindlimb representation. We have investigated the possibility that the reactivation of the deprived hindlimb cortex may be mediated by corticocortical projections from normal to reorganized cortex. The primary somatosensory (SI) cortex was initially mapped to determine the boundaries of the normal and reorganized cortical representations. Somatotopically corresponding regions in both normal and reorganized cortex representing the trunk, the web space, or the shoulder were more precisely mapped. Inactivation of normal cortex was achieved by the nanoinjection of a solution of lidocaine hydrochloride stained with Chicago sky blue. Two major findings are described. First, inactivation of a circumscribed region of normal cortex representing a given receptive field (RF) failed to reduce or inhibit the responsiveness of a somatotopically corresponding RF represented in reorganized cortex. Therefore, it is unlikely that intracortical connections between normal and reorganized cortex could account for the reorganizational processes observed in cats that sustained spinal cord transection at 2 weeks of age. Second, the chemical blockade of normal cortex provoked an increase of the responsiveness and of the size of the peripheral RFs represented in reorganized cortex. This finding suggests that there are corticocortical connections (possibly topographically organized) between normal and reorganized cortex, and that these connections are inhibitory.     

Chronic effects of total or partial digit denervation on raccoon somatosensory cortex.

Electrophysiological recordings were made in the primary somatosensory cortex of anesthetized raccoons 14 to 169 days following digit amputation or 60 to 129 days after transection of the two nerves innervating the ventral surface of the fourth digit. The incidence of inhibitory responses decreased from 50% of the penetrations immediately after amputation to 35% over the first 3 weeks and to almost zero after 2 months. The number of sites with low-threshold excitatory responses increased from 4% to 14% to 50% during these same intervals. Initially, the excitatory fields were small and located over the nerve stumps, and were therefore probably due to direct stimulation of the damaged nerves. At 2 months after amputation, the excitatory receptive fields were large and diffuse. Although the size of receptive fields decreased during the later period (when the thresholds were also decreasing), there was no recovery of any precise somatotopic organization in the deafferented cortex. The reorganization process in the raccoon thus consists of at least two stages: The early stage is dominated by inhibitory connections, whereas the second involves a recovery and restructuring of excitatory inputs. From 2 to 4 months after partial digit denervation, there were only minor changes in response properties or somatotopic organization in the deafferented cortex as compared to immediately after nerve transection. Thus, few of the characteristics of reorganization induced by digit amputation were elicited by this treatment, which leaves some of the digit innervation intact. There was, however, an unexpected increase in the portion of the ventral digit that was able to activate the cortex, suggesting complexities in the peripheral innervation of the digit that need to be resolved.     

Responsiveness of Reorganized Primary Somatosensory (SI) Cortex after Local Inactivation of Normal SI Cortex in Chronic Spinal Cats

The cortical map of adult cats that sustained spinal cord transection at T₁₂ when they were 2 weeks old is characterized by a clear duplication of the representation of the forelimb, rostral trunk, and neck. The novel representation is located in the cortical region that is, in nonoperated animals, normally devoted to the hindlimb representation. We have investigated the possibility that the reactivation of the deprived hindlimb cortex may be mediated by corticocortical projections from normal to reorganized cortex. The primary somatosensory (SI) cortex was initially mapped to determine the boundaries of the normal and reorganized cortical representations. Somatotopically corresponding regions in both normal and reorganized cortex representing the trunk, the web space, or the shoulder were more precisely mapped. Inactivation of normal cortex was achieved by the nanoinjection of a solution of lidocaine hydrochloride stained with Chicago sky blue. Two major findings are described. First, inactivation of a circumscribed region of normal cortex representing a given receptive field (RF) failed to reduce or inhibit the responsiveness of a somatotopically corresponding RF represented in reorganized cortex. Therefore, it is unlikely that intracortical connections between normal and reorganized cortex could account for the reorganizational processes observed in cats that sustained spinal cord transection at 2 weeks of age. Second, the chemical blockade of normal cortex provoked an increase of the responsiveness and of the size of the peripheral RFs represented in reorganized cortex. This finding suggests that there are corticocortical connections (possibly topographically organized) between normal and reorganized cortex, and that these connections are inhibitory.     

Responses in the First or Second Somatosensory Cortical Area in Cats during Transient Inactivation of the Other Ipsilateral Area with Lidocaine Hydrochloride

Simultaneous recordings were obtained from the primary and secondary somatosensory cortical areas (SI and SII) in cats anesthetized with ketamine or pentobarbital. A total of 40 individual neurons were studied (29 in SII and 11 in SI) before, during, and following injections of microliter quantities of lidocaine hydrochloride in the other ipsilateral cortical area. Activity in the cortex injected with the local anesthetic was monitored with single-neuron, multi-neuron, or evoked potential responses to determine the time course of inactivation within 0.5-2 mm of the injection sites. Recording sites in both cortical locations were in the representations of the distal forelimb. Responses were elicited by transcutaneous electrical stimulation across the receptive fields with needle electrodes. Short-latency responses were synchronously activated, and, in those circumstances where single neurons were isolated in both areas, no overall differences in latency were noted. Anesthetization of either cortical area never blocked access of somatosensory information to the intact area, even when the injected cortex was completely silenced in the vicinity of the injection mass. In 15 SII neurons and 7 SI neurons, changes were seen in short-latency evoked responses to stimulation of their receptive fields or in background activity following local anesthesia of the other area through several cycles of injection and recovery. In 7 of these 15 SII cells, changes were noted in the timing and/or firing rates of the short-latency responses; changes were noted in the short-latency responses of 2 of these 7 SI cells while SII was silenced. In 11 SII and 6 SI cells, “background” activity that was recorded during the interstimulus intervals either increased (most cases) or decreased during local anesthesia of the other area. The results are discussed in reference to the hypothesis that primary sensory cortical areas feed information forward to secondary areas, and these feed back modulatory controls to the primary regions.