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PubMed1995

Interaction between afferent input from fingers in human somatosensory cortex.

We recorded somatosensory evoked magnetic fields from eight healthy subjects with a 122-channel whole-scalp SQUID magnetometer. The stimulus sequence consisted of 'standard' stimuli (85%) delivered to palmar side of the left thumb with an interstimulus interval of 0.6 s and of 'deviants' (15%), randomly interspersed among the standards, to little finger, and vice versa. Both stimuli activated four source areas: the contralateral primary somatosensory cortex (SI), the contra-and ipsilateral secondary somatosensory cortices (SII), and the contralateral posterior parietal cortex (PPC). The short-latency (20-40 ms) responses originated in the SI cortex, whereas long-latency responses arose from all 4 areas. At SII and PPC, the deviant stimuli elicited larger responses when presented alone, without intervening standards, than among standards. This implies interaction between afferent impulses from the two fingers and/or partly intermingled cortical representations. Our findings show, in agreement with animal data, different excitatory/inhibitory balance in the various somatosensory areas.

Adult↗
PubMed1987

Functional integration of fetal cortical grafts into the afferent pathway of the rat somatosensory cortex (SmI).

The somatosensory area of fetal (E15-16) Sprague-Dawley rat neocortex was implanted into the barrel receptor area of adult first somatosensory cortex (SmI) to determine if it could be integrated into the host vibrissa-cortical pathway. Functional integration was tested 3-6 months later using the 2-deoxyglucose [44,45] (2-dg) method for estimation of glucose utilization. A variety of control grafts (cerebellum in SmI cortex, neocortex outside the host SmI cortex and appropriately placed neocortex in non-stimulated hosts) all had uniform 2-dg uptake (70 +/- 10 mumol/100 g/min) throughout each graft which averaged 42% less than non-stimulated host SmI cortex (121 +/- 6 mumol/100 g/min; p less than 0.001) but was 3-fold greater than white matter (24.8 +/- 3.1 mumol/100 g/min; p less than 0.001). In appropriately placed neocortical grafts, vibrissal stimulation produced 125% greater average 2-dg uptake (157 +/- 19 mumol/100 g/min) than control grafts (p less than 0.01). Such appropriately placed and stimulated neocortical grafts also contained focal areas of increased 2-dg uptake which were 43% greater than average graft uptake. These data suggest that fetal neocortical grafts were functionally integrated into the physiologically relevant afferent pathways of adult host brains.

Afferent Pathways↗
PubMed1998

Single-cell correlates of a representational boundary in rat somatosensory cortex.

In primary somatosensory cortex (S1), the transition from one representation to the next is typically abrupt when assayed physiologically. However, the extent of anatomical projections to and within the cortex do not strictly respect these physiologically defined transitions. Physiological properties, such as synaptic strengths or intracortical inhibition, have been hypothesized to account for the functionally defined precision of these representational borders. Because these representational borders can be translocated across the cortex by manipulations or behaviors that change the activity patterns of inputs to the cortex, understanding the physiological mechanisms that delimit representations is also an important starting point for understanding cortical plasticity. A novel in vivo and in vitro preparation has been developed to examine the cellular and synaptic mechanisms that underlie representational borders in the rat. In vivo, a short segment of the border between the forepaw-lower jaw representations in rat S1 was mapped using standard electrophysiological methods and was visibly marked using iontophoresis of pontamine sky blue dye. Slices were then obtained from this marked region and maintained in vitro. Intracellularly recorded responses to electrical stimulation of supragranular cortex were obtained from single neurons near the border in response to stimulation within the representational zone or across the border. Both excitatory and inhibitory responses were smaller when evoked by stimuli that activated projections that crossed borders, as compared with stimuli to projections that did not. These findings indicate that intracortical network properties are contributing to the expressions of representational discontinuities in the cortex.

2-Amino-5-phosphonovalerate↗
PubMed2004

Abnormal response recovery in the right somatosensory cortex of dyslexic adults.

Somatosensory evoked fields (SEFs) to repetitive tactile stimuli were recorded from eight dyslexic and eight normal-reading adults. Three successive stimuli, produced by diaphragms driven by compressed air, were delivered to thumb, index finger and thumb in sequence, with stimulus-onset asynchronies (SOAs) of 100 and 200 ms in different runs. Both hands were stimulated alternatingly with an intertrain interval of 1 s, and the responses were recorded with a whole-scalp neuromagnetometer. Whereas the primary somatosensory cortex responses to the first stimuli of the trains did not differ between dyslexics and controls, responses to the second stimuli (and the ratios of second to first responses) were significantly smaller in dyslexic than in control subjects in the right hemisphere (differences 41 and 28% for response amplitudes at the 100 and 200 ms SOAs). The results agree with the proposed pansensory nature of temporal processing deficits in dyslexia, specifically demonstrating abnormal response recovery in the right somatosensory cortex.

Adaptation, Physiological↗
PubMed2000

Differential effects of pain and spatial attention on digit representation in the human primary somatosensory cortex.

Reorganization of primary somatosensory cortex subsequent to either reduced or enhanced peripheral input is well established. Recently, plastic changes following arm amputation in humans were shown to correlate with phantom limb pain. This raised the question whether spatial attention and pain may cause cortical reorganization in the absence of deafferentation. Using non-invasive neuroelectric imaging to study the digit representation in the human primary somatosensory cortex, we report a delayed shift of the representation of digits 2-3 due to pain on the digits 4-5, which outlasted the pain by several minutes. In contrast, reorganization during spatial attention was less pronounced, was seen almost immediately and only during the condition. These data indicate that spatial attention and pain without peripheral deafferentation cause cortical reorganization by different mechanisms. The differential time course of reorganizational effects observed at the cortex may be due to modulation of the lemniscal pathways by nociceptive input from the spinal cord dorsal horn.

Adult↗
PubMed1991

Potential of visual cortex to develop an array of functional units unique to somatosensory cortex.

The identification of specialized areas in the mammalian neocortex, such as the primary visual or somatosensory cortex, is based on distinctions in architectural and functional features. The extent to which certain features that distinguish neocortical areas in rats are prespecified or emerge as a result of epigenetic interactions was investigated. Late embryonic visual cortex transplanted to neonatal somatosensory cortex was later assayed for "barrels," anatomically identified functional units unique to somatosensory cortex, and for boundaries of glycoconjugated molecules associated with barrels. Barrels and boundaries form in transplanted visual cortex and are organized in an array that resembles the pattern in the normal barrelfield. These findings show that different regions of the developing neocortex have similar potentials to differentiate features that distinguish neocortical areas and contribute to their unique functional organizations.

Acetylcholinesterase↗
PubMed2001

Early activation of the primary somatosensory cortex without conscious awareness of somatosensory stimuli in tumor patients.

The primary sensory cortex has usually been regarded as a necessary step in the information processing stream leading to conscious awareness. Recently, it has been proposed that that higher order associative areas rather than the primary sensory areas are the neural basis of conscious perception. In two patients with tumors near the central region we recorded magnetic somatosensory evoked fields. Magnetic source imaging revealed early (40 ms) neural activation in primary somatosensory cortex and absence of later (>60 ms) neural activation in the primary and associative areas in these patients. None of the patients showed conscious awareness of somatosensory stimuli applied to the corresponding body site although the first component of the evoked field was within normal limits. The time course of the magnetic responses and additional evidence on intensity ratings of somatosensory stimuli suggest that early activity in the primary somatosensory cortex is not sufficient for conscious experience to emerge.

Adult↗
PubMed1995

Trigeminal evoked potentials in somatosensory cortex of the Macaca mulatta.

Somatosensory evoked potentials (SEPs) obtained in response to stimulation of the median nerve, posterior tibial nerve, lips, tongue, palate, and pharynx were recorded in four monkeys (Macaca mulatta) under light barbiturate anesthesia. In agreement with the results of SEP recordings in humans and single-unit recordings in monkeys, there is a medial-to-lateral representation in somatosensory cortex of the hand, lips, and tongue. There is a discontinuity in the representation of the upper mouth: the tongue representation is interposed between a medial region (near the lip representation), usually representing the lateral hard palate and gum, and a lateral region (near the lateral sulcus), usually representing the central hard palate. For all types of trigeminal stimulation, root mean square (RMS) voltage maps show maxima over somatosensory cortex. Laminar recordings demonstrated that trigeminal SEPs recorded from the cortical surface are generated in somatosensory cortex in areas 1, 2, and 3b. Median nerve and posterior tibial nerve stimulation did not evoke SEPs in the surface cortex near the lateral sulcus, suggesting that the most lateral portion of the postcentral gyrus is not a part of the second somatosensory area (SII). This lateral region may contain the representation of posterior intraoral structures, but it was not possible to confirm this assumption by stimulation of the soft palate or pharynx.

Animals↗
PubMed2004

Differences between primary somatosensory cortex- and vertex-derived somatosensory-evoked potentials in the rat.

The somatosensory-evoked potential (SEP) elicited by high-intensity stimulation potentially provides a reliable indicator of analgesic efficacy since it reflects the level of activation of the nociceptive system. In the present study, components in the 10-30-ms latency range of SEPs recorded over the primary somatosensory cortex (SI-SEPs) and vertex (Vx-SEP) in the rat were characterized and compared. SEPs were elicited by electrical tail-base stimulation, and SI-SEPs and Vx-SEPs were recorded simultaneously. Responses to increasing stimulus intensity and stimulus frequency while awake and responses to bolus injection of fentanyl, thiopental, and ketamine were investigated. The SI-SEP positive component (P) occurring at 12 ms after stimulation (P12) showed a significantly lower intensity threshold and was significantly less affected by increasing stimulus frequency and by administration of the different drugs when compared to the Vx-SEP P15. The fact that a single stimulus modality results in different signal characteristics dependent on the recording site supports the view that different neural mechanisms involved in primary processing of somatosensory information are responsible for the generation of the SI-SEP P12 and Vx-SEP P15, respectively. This differentiation between SI-SEPs and Vx-SEPs potentially has distinct consequences using the SEP to evaluate nociception and analgesia in the rat model.

Anesthetics↗
PubMed1995

Somatotopy of human hand somatosensory cortex revealed by dipole source analysis of early somatosensory evoked potentials and 3D-NMR tomography.

Somatosensory evoked potentials (SEPs) to median nerve and finger stimulation were analyzed by means of spatio-temporal dipole modelling combined with 3D-NMR tomography in 8 normal subjects. The early SEPs were modelled by 3 equivalent dipoles located in the region of the brain-stem (B) and in the region of the contralateral somatosensory cortex (T and R). Dipole B explained peaks P14 and N18 at the scalp. Dipole T was tangentially oriented and explained the N20-P20, dipole R was radially oriented and modelled the P22. The tangential dipole sources T were located within a distance of 6 mm on the average and all were less than 9 mm from the posterior bank of the central sulcus. In 6 subjects the tangential sources related to finger stimulation arranged along the central sulcus according to the known somatotopy. The radial sources did not show a consistent somatotopic alignment across subjects. We conclude that the combination of dipole source analysis and 3D-NMR tomography is a useful tool for functional localization within the human hand somatosensory cortex.

Adult↗
PubMed1996

Long-lasting potentiation in the secondary somatosensory cortex affects motor control: assessment by H-reflex.

We investigated descending projections from the secondary somatosensory cortex to the feline spinal cord and the effects of long-lasting potentiation in secondary somatosensory cortex on the activities of motoneurons of the cat. Electrophysiological examinations revealed that the low-intensity subthreshold secondary somatosensory cortex stimulation could change the H-Reflex induced by radial nerve stimulation. The H-wave amplitudes, recorded in wrist flexor muscles, were enhanced when the intervals from secondary somatosensory cortex to radial nerve stimuli were altered from 0 to 30 ms (initial excitation, 146 +/- 11% (mean +/- S.E.M.) of the control value). In contrast, the H-waves were suppressed with intervals longer than 30 ms (80 +/- 3%). The descending pathways from secondary somatosensory cortex to the spinal cord were assessed using an immunohistochemical technique. c-Fos and Zif268 proteins, induced by stimulation of the hand-represented secondary somatosensory cortex areas, could thus express in activated cervical neurons. The density of labeled cells was significantly higher in the seventh and eighth cervical segments than in other levels. The great majority of positive cells were distributed in the lateral part of the contralateral ventral horn and their somas ranged from 10 to 50 microns in size. Finally, we examined the effects of long-lasting potentiation, induced by high-frequency stimulation of the ventral posterolateral thalamic nucleus, on the activities of spinal motoneurons. Long-lasting potentiation altered the previously observed effects of secondary somatosensory cortex stimulation on the H-wave amplitude. The secondary somatosensory cortex-conditioned initial excitation of the H-reflex was enhanced (from 139 to 175%, P < 0.05), while late suppression was completely blocked (from 74 to 112%, P < 0.01). In conclusion, the descending pathways from secondary somatosensory cortex to the spinal cord modulated the H-reflex, and long-lasting potentiation in secondary somatosensory cortex affected this modulation. We have previously reported that corticocortical inputs from primary to secondary somatosensory cortex is required for induction of long-lasting potentiation in secondary somatosensory cortex. Taken together, the present study suggests that cortical plasticity in secondary somatosensory cortex amplifies somatic inputs from primary somatosensory cortex as a means of adaptive motor control by the sensory system.

Animals↗
PubMed1992

Population analysis of single neurons in cat somatosensory cortex.

Single neurons in the somatosensory cortex are divisible into a population with receptive fields and a population without receptive fields. These two populations display different laminar distributions, and their respective functions are unknown. We compared other physiological characteristics of these two neuronal populations in an attempt to understand why some neurons lack a receptive field. Only 23% of 465 neurons isolated in the somatosensory cortex of halothane-anesthetized cats displayed a cutaneous receptive field. The iontophoretic administration of glutamate uncovered input from the periphery in another 34% of the sample, leaving 43% of the neurons without evidence of peripheral input under these experimental conditions. Neurons with a receptive field were spontaneously active much more often than neurons lacking peripheral inputs, and their rates of discharge were higher. No differences were found between neurons having a receptive field uncovered with glutamate and those unaffected by glutamate. In all classes of neurons, those cells with spontaneous activity were excited by smaller amounts of glutamate than were silent neurons, but sensitivity to glutamate was not correlated with the presence or absence of a receptive field. We infer that some classes of somatosensory cortical neurons receive strong thalamocortical inputs, whereas others have only relatively weak or no thalamocortical connections. In other experiments we have shown also that those neurons lacking a receptive field and/or spontaneous activity were more likely to be plastic than those with stronger inputs (see Warren and Dykes, 1993a,b), suggesting that neurons having weaker afferent inputs can be more readily modified under certain circumstances.

Afferent Pathways↗
PubMed2006

Involvement of protein kinase A in patterning of the mouse somatosensory cortex.

Patterning of the mouse somatosensory cortex is unusually evident because of the presence of a "barrel field." Presynaptic serotonin and postsynaptic glutamate receptors regulate barrel formation, but little is known of the intracellular signaling pathways through which they act. To determine whether protein kinase A (PKA) plays a role in the development of the barrel field, we examined five viable PKA subunit-specific knock-out (KO) mouse lines for barrel field abnormalities. Barrels are present in these mice, but those lacking the RIIbeta subunit display significantly reduced contrast between the cell densities of barrel hollows and sides compared with wild-type animals. Thalamocortical afferent segregation in the posterior medial barrel subfield appeared normal, suggesting a postsynaptic site of gene action for the RIIbeta protein. Immunoelectron microscopy confirmed that RIIbeta was selectively localized to dendrites and dendritic spines. Mice lacking RIIbeta show reduced glutamate receptor A (GluRA) subunit insertion into the postsynaptic density in postnatal day 7 somatosensory cortex; however, GluRA KO mice developed normal barrels. Our results clearly demonstrate a role for postsynaptic PKA signaling pathways in barrel differentiation. They also demonstrate a clear dissociation between the regulation of GluRA trafficking by PKA and its role in barrel formation. Finally, although a role for PKA downstream of cAMP cannot be ruled out, these data suggest that PKA may not be the principle downstream target because none of the mutants showed a barrelless phenotype similar to that observed in adenylate cyclase type 1 KO mice. These results give insight into activity-dependent mechanisms that regulate barrel formation.

Animals↗
PubMed1997

Histogenesis of ferret somatosensory cortex.

The ferret has emerged as an important animal model for the study of neocortical development. Although detailed studies of the birthdates of neurons populating the ferret visual cortex are available, the birthdates of neurons that reside in somatosensory cortex have not been determined. The current study used bromodeoxyuridine to establish when neurons inhabiting the somatosensory cortex are generated in the ferret; some animals also received injections of [3H]thymidine. In contrast to reports of neurogenesis in ferret visual cortex, most neurons populating the somatosensory cortex have been generated by birth. Although components of all somatosensory cortical layers have been produced at postnatal day 0, the layers are not distinctly formed but develop over a period of several weeks. A small number of neurons continue to be produced for a few days postnatally. The majority of cells belonging to a given layer are born over a period of approximately 3 days, although the subplate and last (layer 2) generated layer take somewhat longer. Although neurogenesis of the neocortex begins along a similar time line for visual and somatosensory cortex, the neurons populating the visual cortex lag substantially during the generation of layer 4, which takes more than 1 week for ferret visual cortex. Layer formation in ferret somatosensory cortex follows many established principles of cortical neurogenesis, such as the well-known inside-out development of cortical layers and the rostro-to-caudal progression of cell birth. In comparison with the development of ferret visual cortex, however, the generation of the somatosensory cortex occurs remarkably early and may reflect distinct differences in mechanisms of development between the two sensory areas.

Animals↗
PubMed1990

Separate finger representations at the human second somatosensory cortex.

We recorded neuromagnetic responses of the second somatosensory cortex in healthy humans. Cutaneous electrical stimulation of fingers elicited a response around 100 ms, with a field pattern agreeing with activation of the second somatosensory cortex in the upper bank of the Sylvian fissure. In an oddball paradigm, with standards presented to the thumb and deviants (10%) to the middle finger, or vice versa, the second somatosensory cortex responses to deviants were almost three times as high in amplitude as those to standards. A similar amplitude enhancement was obtained when the deviants were presented in the absence of the intervening standards but with the same interstimulus interval. The results indicate that an accurate functional representation of different body areas is maintained at the human second somatosensory cortex.

Adult↗
PubMed2006

Current source density analysis of laser heat-evoked intra-cortical field potentials in the primary somatosensory cortex of rats.

The role of the primary somatosensory cortex in thermal pain perception has been established. However, the cortical circuitry that mediates the thermo-nociceptive information processing has not been elucidated. The aim of present study was to investigate the intracortical synaptic currents in primary somatosensory cortex evoked by short laser pulses and to determine their transmission pathway. Noxious CO2 laser pulse stimuli or innocuous electrical and mechanical stimuli were delivered to the hind paw of halothane-anesthetized rats. Multi-channel field potentials were recorded simultaneously in primary somatosensory cortex and laminar-specific transmembrane currents were analyzed using a current source density method. A distinct spatial-temporal pattern of intra-cortical sink source currents was evoked by laser pulse stimuli. The amplitude of the early component was graded by laser energy output and influenced by contralateral signals, whereas the late components were not intensity-dependent and exhibited bilateral excitation. Intra-cortical current flows revealed that synaptic activation occurred initially at layers IV and VI separately and then was relayed transynaptically to the more superficial and the deeper layers. Latency, amplitude and intracortical distributions of the activated intra-cortical currents evoked by noxious stimuli differed significantly from those evoked by innocuous stimuli. Conduction velocity data together with the results of tetrodotoxin, capsaicin and morphine treatments indicated that the early and late components were mediated separately by A-delta and C fibers. Our results suggest that large and small diameter thermal nociceptive afferents generated laminar-specific intracortical synaptic currents in primary somatosensory cortex and that these excitatory synaptic currents were conveyed separately by lateral and medial thalamic nuclei.

Animals↗
PubMed2005

Large-scale changes in dendritic structure during reorganization of adult somatosensory cortex.

In adult rat somatosensory cortex (S1), neurons are biased and have less dendritic arbor close to the border between the forepaw and lower jaw representations. Changes in sensory experience cause changes in the functional organization of the neocortex. Therefore, we examined the morphology of neurons in the reorganized region of S1 after forepaw denervation. We found that during reorganization dendritic arbors changed to reflect the new location of the border.

Animals↗
PubMed2000

Distribution of neurons immunoreactive for calcium-binding proteins varies across areas of cat primary somatosensory cortex.

The primary somatosensory (SI) cortex in the cat contains four cytoarchitectonic areas that appear to contain separate body representations and have different functions. We tested whether functional differences among these areas are reflected in the densities of neurons containing each of three calcium-binding proteins: parvalbumin (PV), calbindin (CB), and calretinin (CR). Colocalization experiments revealed that CR was localized in a population of neurons distinct from those containing PV or CB. The general laminar distributions of the three calcium-binding proteins were similar to those described in other species and cortical areas, but there were significant density differences in layers II and III across SI. The density of PV-immunoreactive neurons was higher in areas 3b and 1 than in areas 3a and 2. CB-immunoreactive neurons were found in higher densities in anterior SI than in posterior SI, and the pattern of CR-immunoreactive neurons was reciprocal to that of CB, with significantly higher densities in posterior regions of SI. Since the firing characteristics of nonpyramidal neurons appear to be related to their calcium-binding protein content, differences in regional distributions of these neurons in layers II and III may contribute to functional differences between the cytoarchitectonic areas of SI cortex.

Animals↗