Spatiotemporal dynamics of bimanual integration in human somatosensory cortex and their relevance to bimanual object manipulation

Little is known about the spatiotemporal dynamics of cortical responses that integrate slightly asynchronous somatosensory inputs from both hands. This study aimed to clarify the timing and magnitude of interhemispheric interactions during early integration of bimanual somatosensory information in different somatosensory regions and their relevance for bimanual object manipulation and exploration. Using multi-fiber probabilistic diffusion tractography and MEG source analysis of conditioning-test (C-T) median nerve somatosensory evoked fields in healthy human subjects, we sought to extract measures of structural and effective callosal connectivity between different somatosensory cortical regions and correlated them with bimanual tactile task performance. Neuromagnetic responses were found in major somatosensory regions, i.e., primary somatosensory cortex SI, secondary somatosensory cortex SII, posterior parietal cortex, and premotor cortex. Contralateral to the test stimulus, SII activity was maximally suppressed by 51% at C-T intervals of 40 and 60 ms. This interhemispheric inhibition of the contralateral SII source activity correlated directly and topographically specifically with the fractional anisotropy of callosal fibers interconnecting SII. Thus, the putative pathway that mediated inhibitory interhemispheric interactions in SII was a transcallosal route from ipsilateral to contralateral SII. Moreover, interhemispheric inhibition of SII source activity correlated directly with bimanual tactile task performance. These findings were exclusive to SII. Our data suggest that early interhemispheric somatosensory integration primarily occurs in SII, is mediated by callosal fibers that interconnect homologous SII areas, and has behavioral importance for bimanual object manipulation and exploration.     

The organization of thalamocortical relay neurons in the rat ventrobasal complex studied by the retrograde transport of horseradish peroxidase.

The organization of thalamocortical relay neurons in the thalamic ventrobasal complex (VB) of the rat was investigated by the use of the retrograde axonal transport of horseradish peroxidase (HRP). Injections of HRP into somatosensory cortex (SI) resulted in a distinctive gradient of neuronal and non-neuronal HRP reaction product. Electrophysiologically characterized points of SI injected with small volumes of HRP labeled a sector of neurons in VB ipsilateral to the injection. This zone of labeled neurons consisted of a complex curvilinear array or lamina with a rostral hollow or solid expansion of densely packed HRP positive neurons which continues caudally as a less dense tapering wing. Despite this complex arrangement, an approximate pattern of somatotopy was determined indicating the direction of shifts in peripheral receptive fields moving in any axis of VB. The number of labeled neurons projecting upon a point in SI was also determined. Injections of the cortical vibrissae, face and forepaw representations labeled a greater number of neurons in VB per unit area of cortex than did injections in the hindlimb or body representations. The total number of HRP positive neurons in VB increased proportionately with areal increase of the HRP injections into overlapping cortical representations. The number of HRP positive neurons in the rostral half of the laminae increased almost linearly with the area of injection, while the number of HRP positive neurons in the caudal half of the laminae showed relatively smaller increases.     

Altered somatosensory processing in trigeminal neuralgia

Trigeminal neuralgia (TN) is a pain state characterized by intermittent unilateral pain attacks in one or several facial areas innervated by the trigeminal nerve. The somatosensory cortex is heavily involved in the perception of sensory features of pain, but it is also the primary target for thalamic input of nonpainful somatosensory information. Thus, pain and somatosensory processing are accomplished in overlapping cortical structures raising the question whether pain states are associated with alteration of somatosensory function itself. To test this hypothesis, we used functional magnetic resonance imaging to assess activation of primary (SI) and secondary (SII) somatosensory cortices upon nonpainful tactile stimulation of lips and fingers in 18 patients with TN and 10 patients with TN relieved from pain after successful neurosurgical intervention in comparison with 13 healthy subjects. We found that SI and SII activations in patients did neither depend on the affected side of TN nor differ between operated and nonoperated patients. However, SI and SII activations, but not thalamic activations, were significantly reduced in patients as compared to controls. These differences were most prominent for finger stimulation, an area not associated with TN. For lip stimulation SI and SII activations were reduced in patients with TN on the contra‐ but not on the ipsilateral side to the stimulus. These findings suggest a general reduction of SI and SII processing in patients with TN, indicating a long‐term modulation of somatosensory function and pointing to an attempt of cortical adaptation to potentially painful stimuli. Hum Brain Mapp, 2009. © 2009 Wiley‐Liss, Inc.     

Corticothalamic and corticotectal somatosensory projections from the anterior ectosylvian sulcus (SIV cortex) in neonatal cats: an anatomical demonstration with HRP and 3H-leucine

Corticothalamic and corticotectal projections from the anterior ectosylvian sulcus (AES) in neonatal cats were studied with anterograde and retrograde neuroanatomical techniques. When the injection site was relatively restricted to the sulcal walls and fundus of the rostral AES (i.e., the SIV cortex), heavy ipsilateral thalamic label was observed in the medial subdivision of the posterior group, in the suprageniculate nucleus, and in the external medullary lamina. No terminal label was seen in the contralateral thalamus although the contralateral homotopic cortex was heavily labeled. Within the ventrobasal complex (VB), dense axonal label was observed in fascicles that traversed VB, but only light terminal label was observed within VB itself. However, in cases where the tracer spread into adjacent SII, terminal label in VB was pronounced. Similarly, when the injection site extended into auditory cortex, terminal label was observed in the lateral and intermediate subdivisions of the posterior group. Rostral AES injections produced distinct, predominantly ipsilateral, terminal label in the superior colliculus that was distributed in two tiers: a discontinuous band in the stratum griseum intermedium and a more diffuse band in stratum griseum profundum. Caudally, dense terminal label was seen in the intercollicular zone and dorsolateral periaqueductal gray. When the injection site did not include rostral AES, no label was observed in the superior colliculus. Horseradish peroxidase injections into the superior colliculus of neonates produced retrogradely labeled neurons throughout the AES, but none was found on the crown of the gyrus where SII is located. Thus, the neonatal corticotectal somatosensory projection arises exclusively from AES and parallels that found in adults. These data indicate that the elaboration of a major descending somatosensory pathway from AES to the thalamus and midbrain is largely a prenatal event. The in utero anatomical maturation of the corticofugal projections from SIV cortex to the superior colliculus contrasts with the protracted postnatal development of the corticotrigeminal projections from SI cortex but is consistent with the mature anatomical state of ascending trigeminotectal projections.     

Corticofugal influences of S1 cortex on ventrobasal thalamic neurons in the awake rat

Corticofugal influences on the responses of 39 ventrobasal (VB) thalamic neurons to repetitive stimuli were studied in awake rats by focally suppressing the evoked and spontaneous electrocortical activity of the primary (S1) somatosensory cortex with magnesium or lidocaine plus magnesium. Suppression of the S1 cortex reduced the number of spikes discharged by 19 (66%) of 29 VB units in response to each of 25 electrical stimuli delivered to the medial lemniscus; 9 units were unaffected and 1 showed an increased response to 1-10 Hz stimuli. The responses of 6 (38%) of 16 VB units to electrical somatic stimuli were also reduced following S1 cortical suppression; 9 units were unaffected and 1 showed an increased response to 20-40 Hz stimuli. A comparison of the reduced responsiveness of 5 units studied during medial lemniscal and somatic stimulation did not reveal any additional response attenuation attributable to subthalamic corticofugal influences. We conclude that, in the awake rat, somatosensory transmission to VB thalamic neurons is primarily facilitated by S1 corticothalamic neurons.     

Overlapping corticostriatal projections from the rodent vibrissal representations in primary and secondary somatosensory cortex

To determine whether the neostriatum receives overlapping projections from two somatosensory cortical areas, the anterograde tracers biotinylated dextran amine (BDA) and fluoro-ruby (FR) were injected into the whisker representations of primary (SI) and secondary (SII) somatosensory cortex. Reconstructions of labeled terminals and their beaded varicosities in the neostriatum and thalamus were analyzed quantitatively to compare the extent of overlapping projections to both subcortical structures. Corticostriatal projections from focal sites in both somatosensory areas exhibited substantial amounts of divergence within the dorsolateral neostriatum. Most of the labeled terminals were concentrated in densely packed arborizations that occupied lamellar-shaped regions along the dorsolateral edge of the neostriatum. Tracer injections in both cortical areas also produced dense anterograde and retrograde labeling in the thalamus, especially in the ventrobasal complex (VB) and in the medial part of the posterior (POm) nucleus. Because these thalamic regions are topographically organized and have reciprocal connections with corresponding representations in both SI and SII, the amount of labeled overlap in the thalamus was used to indicate the degree of somatotopic correspondence at the SI and SII injection sites. We found that the proportion of overlapping projections to the neostriatum was moderately correlated with the amount of overlap observed in the thalamus. This result strongly indicates that specific sites in the dorsolateral neostriatum receive convergent projections from corresponding somatotopic representations in SI and SII, but also suggests that some of the corticostriatal divergence may reflect neostriatal integration of somatosensory information from noncorresponding representations in SI and SII.     

Age-related changes across the primary and secondary somatosensory areas: An analysis of neuromagnetic oscillatory activities

ObjectiveAge-related changes are well documented in the primary somatosensory cortex (SI). Based on previous somatosensory evoked potential studies, the amplitude of N20 typically increases with age probably due to cortical disinhibition. However, less is known about age-related change in the secondary somatosensory cortex (SII). The current study quantified age-related changes across SI and SII mainly based on oscillatory activity indices measured with magnetoencephalography.MethodsWe recorded somatosensory evoked magnetic fields (SEFs) to right median nerve stimulation in healthy young and old subjects and assessed major SEF components. Then, we evaluated the phase-locking factor (PLF) for local field synchrony on neural oscillations and the weighted phase-lag index (wPLI) for cortico-cortical synchrony between SI and SII.ResultsPLF was significantly increased in SI along with the increased amplitude of N20m in the old subjects. PLF was also increased in SII associated with a shortened peak latency of SEFs. wPLI analysis revealed the increased coherent activity between SI and SII.ConclusionsOur results suggest that the functional coupling between SI and SII is influenced by the cortical disinhibition due to normal aging.SignificanceWe provide the first electrophysiological evidence for age-related changes in oscillatory neural activities across the somatosensory areas.     

Overlapping corticostriatal projections from the rodent vibrissal representations in primary and secondary somatosensory cortex

To determine whether the neostriatum receives overlapping projections from two somatosensory cortical areas, the anterograde tracers biotinylated dextran amine (BDA) and fluoro-ruby (FR) were injected into the whisker representations of primary (SI) and secondary (SII) somatosensory cortex. Reconstructions of labeled terminals and their beaded varicosities in the neostriatum and thalamus were analyzed quantitatively to compare the extent of overlapping projections to both subcortical structures. Corticostriatal projections from focal sites in both somatosensory areas exhibited substantial amounts of divergence within the dorsolateral neostriatum. Most of the labeled terminals were concentrated in densely packed arborizations that occupied lamellar-shaped regions along the dorsolateral edge of the neostriatum. Tracer injections in both cortical areas also produced dense anterograde and retrograde labeling in the thalamus, especially in the ventrobasal complex (VB) and in the medial part of the posterior (POm) nucleus. Because these thalamic regions are topographically organized and have reciprocal connections with corresponding representations in both SI and SII, the amount of labeled overlap in the thalamus was used to indicate the degree of somatotopic correspondence at the SI and SII injection sites. We found that the proportion of overlapping projections to the neostriatum was moderately correlated with the amount of overlap observed in the thalamus. This result strongly indicates that specific sites in the dorsolateral neostriatum receive convergent projections from corresponding somatotopic representations in SI and SII, but also suggests that some of the corticostriatal divergence may reflect neostriatal integration of somatosensory information from noncorresponding representations in SI and SII.     

Characteristics of the human contra- versus ipsilateral SII cortex.

OBJECTIVES: In order to study the interaction between left- and right-sided stimuli on the activation of cortical somatosensory areas, we recorded somatosensory evoked magnetic fields (SEFs) from 8 healthy subjects with a 122 channel whole-scalp SQUID gradiometer. METHODS: Right and left median nerves were stimulated either alternately within the same run, with interstimulus intervals (ISIs) of 1.5 and 3 s, or separately in different runs with a 3 s ISI. In all conditions 4 cortical source areas were activated: the contralateral primary somatosensory cortex (SI), the contra- and ipsilateral secondary somatosensory cortices (SII) and the contralateral posterior parietal cortex (PPC). RESULTS: The earliest activity starting at 20 ms was generated solely in the SI cortex, whereas longer-latency activity was detected from all 4 source areas. The mean peak latencies for SII responses were 86-96 ms for contralateral and 94-97 ms for ipsilateral stimuli. However, the activation of right and left SII areas started at 61+/-3 and 62+/-3 ms to contralateral stimuli and at 66+/-2 and 63+/-2 ms to ipsilateral stimuli, suggesting a simultaneous commencing of activation of the SII areas. PPC sources were activated between 70 and 110 ms in different subjects. The 1.5 s ISI alternating stimuli elicited smaller SII responses than the 3 s ISI non-alternating stimuli, suggesting that a considerable part of the neural population in SII responds both to contra- and ipsilateral stimuli. The earliest SI responses did not differ between the two conditions. There were no significant differences in source locations of SII responses to ipsi- and contralateral stimuli in either hemisphere. Subaverages of the responses in sets of 30 responses revealed that amplitudes of the SII responses gradually attenuated during repetitive stimulation, whereas the amplitudes of the SI responses were not changed. CONCLUSIONS: The present results implicate that ipsi- and contralateral SII receive simultaneous input, and that a large part of SII neurons responds both to contra- and ipsilateral stimulation. The present data also highlight the different behavior of SI and SII cortices to repetitive stimuli.     

Hypothermia-induced changes of afferent sensory transmission to the VPM thalamus of rats and hamsters

Effects of hypothermia on the afferent somatosensory transmission to the ventroposteromedial (VPM) thalamus were determined in anesthetized rats and hamsters. Hamsters showed a gradual suppression of afferent sensory transmission during cooling (to 18 degrees C) and disinhibition during subsequent warming of body temperature (Tb). However, rats exhibited steep inhibition from Tb 26 degrees C to complete absence of sensory transmission at Tb 20 degrees C and abrupt disinhibition during subsequent warming. Species difference at thalamic level was quite similar to our previous results in the primary somatosensory (SI) cortex, suggesting that changes of sensory transmission observed in the SI cortex may have already occurred at thalamic level. Differences between the cortex and the thalamus were observed only during deep hypothermia in rat and during the final period of warming in hamster. Conduction latencies of thalamocortical system of both species were not influenced during Tb lowering until 24 degrees C (equivalent to brain temperature 25-26 degrees C). These results suggest inherently different adaptability to hypothermia in processing somatosensory information between hibernator and non-hibernator, but similar sustainability of sensory functions of the thalamocortical system during hypothermia in both species.     

Adaptive changes in the neuromagnetic response of the primary and association somatosensory areas following repetitive tactile hand stimulation in humans

Cortical adaptation in the primary somatosensory cortex (SI) has been probed using different stimulation modalities and recording techniques, in both human and animal studies. In contrast, considerably less knowledge has been gained about the adaptation profiles in other areas of the cortical somatosensory network. Using magnetoencephalography (MEG), we examined the patterns of short‐term adaptation for evoked responses in SI and somatosensory association areas during tactile stimulation applied to the glabrous skin of the hand. Cutaneous stimuli were delivered as trains of serial pulses with a constant frequency of 2 Hz and 4 Hz in separate runs, and a constant inter‐train interval of 5 s. The unilateral stimuli elicited transient responses to the serial pulses in the train, with several response components that were separated by independent component analysis. Subsequent source reconstruction techniques identified regional generators in the contralateral SI and somatosensory association areas in the posterior parietal cortex (PPC). Activity in the bilateral secondary somatosensory cortex (i.e., SII/PV) was also identified, although less consistently across subjects. The dynamics of the evoked activity in each area and the frequency‐dependent adaptation effects were assessed from the changes in the relative amplitude of serial responses in each train. We show that the adaptation profiles in SI and PPC areas can be quantitatively characterized from neuromagnetic recordings using tactile stimulation, with the sensitivity to repetitive stimulation increasing from SI to PPC. A similar approach for SII/PV has proven less straightforward, potentially due to the tendency of these areas to respond selectively to certain stimuli. Hum Brain Mapp, 2013. © 2012 Wiley Perodicals, Inc.     

Long-lasting potentiation of field potentials in primary and secondary somatosensory cortex

Effects of tetanic bursts (200 Hz, 10 pulses) on field potentials elicited by ventral posterolateral thalamic nucleus (VPL) stimulation were investigated in the feline somatosensory cortex. In the first experiments, field potentials elicited by VPL stimulation (test pulse) were simultaneously recorded in the primary (SI) and the secondary (SII) somatosensory cortex in six animals. Potentiation of field potentials recorded in SII was induced by tetanic stimulation of VPL in all six animals, whereas the same tetanic bursts failed to produce significant changes in SI in four of the six animals. The results suggest that plastic changes in somatosensory processing take place in SII rather than SI. In subsequent experiments, features of the potentiation observed in SII were examined in 20 animals. The field potentials were simultaneously recorded at 16 points placed vertically at 150-μm intervals from the cortical surface. The potentiation of field potentials (to 110–170% of control values) observed at depths between 600 and 1350 μm lasted more than 90 min after tetanic stimulation. Poststimulus histograms of multiple-unit activities revealed a long-lasting increase in the number of unit discharges evoked by VPL stimulation. This change in the number of activated cellsis regarded as a cause of potentiation of SII field potentials. In the last session, the effects of N-methyl-d-aspartate (NMDA) receptor antagonists on the potentiation of SII field potentials were investigated. Cortical intraventricular injection ofd-2-amino-5-phosphonovalerate (APV) anddl-2-amino-7-phosphonoheptanoic acid (APH) prevented induction of the potentiation in SII. NMDA receptor activation participates in forming this SII potentiation.     

A multisensory zone in rat parietotemporal cortex: intra- and extracellular physiology and thalamocortical connections

Multisensory integration is essential for the expression of complex behaviors in humans and animals. However, few studies have investigated the neural sites where multisensory integration may occur. Therefore, we used electrophysiology and retrograde labeling to study a region of the rat parietotemporal cortex that responds uniquely to auditory and somatosensory multisensory stimulation. This multisensory responsiveness suggests a functional organization resembling multisensory association cortex in cats and primates. Extracellular multielectrode surface mapping defined a region between auditory and somatosensory cortex where responses to combined auditory/somatosensory stimulation were larger in amplitude and earlier in latency than responses to either stimulus alone. Moreover, multisensory responses were nonlinear and differed from the summed unimodal responses. Intracellular recording found almost exclusively multisensory cells that responded to both unisensory and multisensory stimulation with excitatory postsynaptic potentials (EPSPs) and/or action potentials, conclusively defining a multisensory zone (MZ). In addition, intracellular responses were similar to extracellular recordings, with larger and earlier EPSPs evoked by multisensory stimulation, and interactions suggesting nonlinear postsynaptic summation to combined stimuli. Thalamic input to MZ from unimodal auditory and somatosensory thalamic relay nuclei and from multisensory thalamic regions support the idea that parallel thalamocortical projections may drive multisensory functions as strongly as corticocortical projections. Whereas the MZ integrates uni- and multisensory thalamocortical afferent streams, it may ultimately influence brainstem multisensory structures such as the superior colliculus.     

Enlarged SI and SII somatosensory evoked responses in the CLN5 form of neuronal ceroid lipofuscinosis.

OBJECTIVES: To examine in detail the activation of the primary (SI) and secondary (SII) somatosensory cortex in CLN5, the Finnish variant of late infantile neuronal ceroid lipofuscinoses (NCL). METHODS: Somatory evoked magnetic fields were recorded with a 122-channel planar gradiometer in response to median nerve stimulation in 5 CLN5 patients (aged 8.8-16.7 years) and in 10 healthy age-matched controls. RESULTS: The first two responses from contralateral SI, N20m and P35m, were 6-20 times stronger in the patients than in the controls. The morphology of the subsequent deflections from SI was abnormal in the patients: a prominent N45m was detected, while the normally present P60m deflection was missing. In 4 patients the contra- and in two patients the ipsilateral SII responses were also enlarged. Furthermore, the SII activation was detected at shorter latency in patients than in controls. CONCLUSIONS: At SI, CLN5 is associated with a selective enhancement of the early cortical responses. We propose that the enlargement of N20m most likely reflects increased synchronous input from thalamus, whereas the altered morphology of the following responses may reflect defective interneuronal inhibition at the cortex. The enlargement of SII responses shows that the imbalance between excitation and inhibition in CLN5 extends outside the primary somatosensory areas.     

Reactions of neurons in the primary and secondary somatosensory zones of the cortex to stimulation of the ventral posterior nucleus of the thalamus]

Neuronal responses in the first and second somatosensory cortex (SI and SII) to stimulation of the ventroposterior nucleus of the thalamus (VP) were studied in experiments on cats immobilized with d-tubocurarine. 12.0% responding neurons in SI and 9.5% in SII were activated antidromically by VP stimulation. In the majority of antidromic responses the latencies did not exceed 1.0 ms. The minimal latency of orthodromic spikes was 1.5 ms in SI and 1.7 ms in SII. In SI the number of neurons whose orthodromic spike latencies did not exceed 3.0 ms was larger than neurons activated with latencies of 3.1-4.5 ms. In SII an inverse quantitative relationship between those two neuronal groups was observed. In SII a significantly larger number of neurons was excited with latencies of EPSPs ranged between 1.1-9.0 ms in SI and between 1.4-6.6 ms in SII and the latencies of IPSPs between 1.5-6.8 ms in SI and 2.2-9.4 ms in SII. The importance of different pathways for excitatory and inhibitory VP influences to neurons of SI and SII is discussed.     

Thalamocortical dysfunction and thalamic injury after asphyxial cardiac arrest in developing rats

Global hypoxia-ischemia interrupts oxygen delivery and blood flow to the entire brain. Previous studies of global brain hypoxia-ischemia have primarily focused on injury to the cerebral cortex and to the hippocampus. Susceptible neuronal populations also include inhibitory neurons in the thalamic reticular nucleus. We therefore investigated the impact of global brain hypoxia-ischemia on the thalamic circuit function in the somatosensory system of young rats. We used single neuron recordings and controlled whisker deflections to examine responses of thalamocortical neurons to sensory stimulation in rat survivors of 9 min of asphyxial cardiac arrest incurred on postnatal day 17. We found that 48-72 h after cardiac arrest, thalamocortical neurons demonstrate significantly elevated firing rates both during spontaneous activity and in response to whisker deflections. The elevated evoked firing rates persist for at least 6-8 weeks after injury. Despite the overall increase in firing, by 6 weeks, thalamocortical neurons display degraded receptive fields, with decreased responses to adjacent whiskers. Nine minutes of asphyxial cardiac arrest was associated with extensive degeneration of neurites in the somatosensory nucleus as well as activation of microglia in the reticular nucleus. Global brain hypoxia-ischemia during cardiac arrest has a long-term impact on processing and transfer of sensory information by thalamic circuitry. Thalamic circuitry and normalization of its function may represent a distinct therapeutic target after cardiac arrest.     

Responsiveness of ventrobasal thalamic neurons after suppression of S1 cortex in the anesthetized rat

Corticofugal influences on the responses of ventrobasal (VB) thalamic neurons to repetitive stimuli were studied in anesthetized rats by suppressing primary somatosensory (S1) electrocortical activity with topically applied lidocaine. Effective concentrations of lidocaine were confined to S1 and immediately adjacent cortex and suppressed evoked S1 responses and corticofugal discharges. Suppression of S1 cortex reduced the average number of spikes discharged by 83 VB neurons in response to each of 25 electrical somatic stimuli delivered at frequencies ranging from 1 to 50 Hz. Of 20 units studied both before and after S1 suppression, 14 (70%) showed a similar reduced response to repetitive stimuli. Cortical suppression produced no consistent changes in spontaneous activity, somatic stimulus threshold, response latency, or size of receptive field. There was no significant difference in the effect of cortical suppression on the responsiveness of 8 VB neurons to repetitive medial lemniscal, as compared to somatic, stimuli. We conclude that, in the anesthetized rat, S1 corticofugal activity facilitates somato-sensory transmission to VB neurons and that this facilitation is mediated, at least in part, by corticothalamic neurons.     

Different cortical organization of visceral and somatic sensation in humans

Sensory stimuli from the visceral domain exhibit perceptual characteristics different from stimuli applied to the body surface. Compared with somatosensation there is not much known about the cortical projection and functional organization of visceral sensation in humans. In this study, we determined the cortical areas activated by non-painful electrical stimulation of visceral afferents in the distal oesophagus, and somatosensory afferents in the median nerve and the lip in seven healthy volunteers using whole-head magnetoencephalography. Stimulation of somatosensory afferents elicited short-latency responses (≈ 20–60 ms) in the primary somatosensory cortex (SI) contralateral (median nerve) or bilateral (lip) to the stimulated side, and long-latency responses (≈ 60–160 ms) bilaterally in the second somatosensory cortex (SII). In contrast, stimulation of visceral oesophageal afferents did not evoke discernible responses in SI but well reproducible bilateral SII responses (≈ 70–190 ms) in close vicinity to long-latency SII responses following median nerve and lip stimuli. Psychophysically, temporal discrimination of successive stimuli became worse with increasing stimulus repetition rates (0.25 Hz, 0.5 Hz, 1 Hz, 2 Hz) only for visceral oesophageal, but not for somatosensory median nerve stimuli. Correspondingly, amplitudes of the first cortical response to oesophageal stimulation emerging in the SII cortex declined with increasing stimulus repetition rates whereas the earliest cortical response elicited by median nerve stimuli (20 ms SI response) remained unaffected by the stimulus frequency. Our results indicate that visceral afferents from the oesophagus primarily project to the SII cortex and, unlike somatosensory afferents, lack a significant SI representation. We propose that this cortical projection pattern forms the neurophysiological basis of the low temporal and spatial resolution of conscious visceral sensation.     

Spinothalamocortical projections to the secondary somatosensory cortex (SII) in squirrel monkey

Anterograde labeling of the cervical spinothalamic tract was combined with retrograde labeling of thalamocortical cells projecting to the hand region of the second somatosensory cortex (hSII) to identify likely sites in the thalamus for processing and transmitting nociceptive information to hSII. Anterograde labeling of terminals was done with 2% WGA-HRP injections in the cervical enlargement; thalamocortical cells were retrogradely labeled with fluorescent tracers. In one experiment, the contralateral primary somatosensory cortex hand region (hSI) was injected to provide a direct comparison with hSII thalamic label. Both labeled cells and terminal-like structures were visualized in single thalamic sections and their numbers and positions quantitatively analyzed. The number of labeled cells within 100 microns from the STT terminals were counted as overlapping cells. Four thalamic nuclei, ventroposterior inferior (VPI), ventroposterior lateral (VPL), posterior nucleus (PO) and centrolateral nucleus (CL) combined to contain 86.5% of all hSII-projecting overlapping cells. Of all hSII-projecting thalamic overlapping cells, VPI contained the largest number (36.4% of the total) followed by the anterior portion of the posterior nuclear complex (POa; 20.4%), VPL (18.3%) and CL (11.4%). Results of the hSI injection show a different pattern of overlap in agreement with our earlier study. The relative distribution of overlapping cells was dependent on the antero-posterior position of the SII injections. The most anterior injections resulted in small numbers of labeled cells, with the majority of overlapping cells located in PO and CL. The more posterior injections resulted in overlapping cells mainly in VPI and VPL. The results indicate that, in the squirrel monkey, VPI, VPL, POa and CL relay nociceptive information from the spinal cord to the second somatosensory cortex.     

Neuromagnetic gamma-band activity in the primary and secondary somatosensory areas

To evaluate the gamma-band activity related to somatosensory processing, we recorded neuromagnetic signals from seven healthy subjects. The source power changes evoked by electrical stimulation of the median nerve were estimated with synthetic aperture magnetometry (SAM). Source power in the low gamma band (40 Hz) decreased in the contralateral primary somatosensory cortex (SI) for a few hundred milliseconds (i.e. middle and long latency) and then increased inversely. Source power in the high gamma band (70-90 Hz) increased simultaneously both in the contralateral SI and contra/ipsilateral secondary somatosensory cortex (SII) in 80-180 ms. These results suggest that low and high gamma oscillations work under independent mechanisms during somatosensory processing. In particular, high gamma oscillations may play an essential role in making a functional connection between SI and SII.