Different laterality between the thumb and index finger in human SII activities

We compared the ipsilateral and contralateral evoked activities in the secondary somatosensory cortex (SII) of four human subjects. Magnetic cortical fields, evoked by electrical stimulation of the distal interphalangeal joint of either the thumb or the index finger, were recorded by an 80-channel whole head type magnetoencephalography system (MEG). We calculated the ratio of the equivalent current dipole (ECD) moment of the ipsilateral area to that of the contralateral area. The mean ratio for the thumb across the subjects was 0.43 +/- 0.13, while that for the index finger was 0.77 +/- 0.20 (mean +/- s.d). The ratio for the index finger was significantly higher than that for the thumb (p = 0.039, two tailed t-test). These results show that the cortical index finger area is activated more bilaterally than the thumb area.     

Primary somatosensory cortex is actively involved in pain processing in human

We recorded somatosensory evoked magnetic fields (SEFs) by a whole head magnetometer to elucidate cortical receptive areas involved in pain processing, focusing on the primary somatosensory cortex (SI), following painful CO(2) laser stimulation of the dorsum of the left hand in 12 healthy human subjects. In seven subjects, three spatially segregated cortical areas (contralateral SI and bilateral second (SII) somatosensory cortices) were simultaneously activated at around 210 ms after the stimulus, suggesting parallel processing of pain information in SI and SII. Equivalent current dipole (ECD) in SI pointed anteriorly in three subjects whereas posteriorly in the remaining four. We also recorded SEFs following electric stimulation of the left median nerve at wrist in three subjects. ECD of CO(2) laser stimulation was located medial-superior to that of electric stimulation in all three subjects. In addition, by direct recording of somatosensory evoked potentials (SEPs) from peri-Rolandic cortex by subdural electrodes in an epilepsy patient, we identified a response to the laser stimulation over the contralateral SI with the peak latency of 220 ms. Its distribution was similar to, but slightly wider than, that of P25 of electric SEPs. Taken together, it is postulated that the pain impulse is received in the crown of the postcentral gyrus in human.     

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.     

Differential modulation in human primary and secondary somatosensory cortices during the preparatory period of self-initiated finger movement

To elucidate the mechanisms underlying sensorimotor integration, we investigated modulation in the primary (SI) and secondary (SII) somatosensory cortices during the preparatory period of a self-initiated finger extension. Electrical stimulation of the right median nerve was applied continuously, while the subjects performed a self-initiated finger extension and were instructed not to pay attention to the stimulation. The preparatory period was divided into five sub-periods from the onset of the electromyogram to 3000 ms before movement and the magnetoencephalogram signals following stimulation in each sub-period were averaged. Multiple source analysis indicated that the equivalent current dipoles (ECDs) were located in SI and bilateral SII. Although the ECD moment for N 20 m (the upward deflection peaking at around 20 ms) was not significantly changed, that for P 30 m (the downward deflection peaking at around 30 m) was significantly smaller in the 0- to -500-ms sub-period than the -2000- to -3000-ms sub-period. As for SII, the ECD moment for the SII ipsilateral to movement showed no significant change, while that for the contralateral SII was significantly larger in the 0- to -500-ms sub-period than the -1500- to -2000-ms or -2000- to -3000-ms sub-period. The opposite effects of movement on SI and SII cortices indicated that these cortical areas play a different role in the function of the sensorimotor integration and are affected differently by the centrifugal process.     

Effects of an acute D2-dopaminergic blockade on the somatosensory cortical responses in healthy humans: evidence from evoked magnetic fields

We tested the possible role of dopaminergic activity in the processing of somatosensory afferent information in healthy humans. Somatosensory evoked magnetic fields (SEFs) were recorded in seven subjects in response to left median nerve stimulation. SEFs were obtained in all subjects after oral administration of 2 mg haloperidol, an antagonist to dopaminergic D2 receptors, and placebo, which were given in a randomized, double-blind cross-over design. SEFs were analyzed using a multiple equivalent current dipole (ECD) model, with one dipole at the right primary somatosensory cortex (SI) and at both left and right secondary somatosensory cortices (SII). The earliest responses from SI, peaking at about 20 ms (N20m) and 35 ms (P35m), were not affected by haloperidol. A later deflection peaking at about 75 ms (P60m), however, was slightly reduced (p < 0.05). Responses arising from SII were not significantly changed. The results suggest that dopaminergic activity may be involved in modulating somatosensory processing after the initial stages of cortical activation.     

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.     

Effects on median nerve SEPs of tactile stimulation applied to adjacent and remote areas of the body surface

Study of the influence of continuous tactile stimulation on somatosensory evoked potentials (SEPs) following electrical stimulation of the median nerve revealed an effect due to interfering input from both adjacent and remote regions of the body surface. The distribution of the effect was demonstrated by subtracting the 'interference' from the 'control' response to derive a 'difference' wave form. Tactile stimulation of the thumb ipsilateral to the stimulated median nerve produced a difference wave form in which a marked phase reversal was apparent between pre- and post-central areas for 2 complexes, at latencies of approximately 20 and 30 msec. It is proposed that this may have been due to partial 'saturation' of a generator in the hand region of area 3b in the primary somatosensory cortex (SI), which was then unable to respond fully to the median nerve impulse. A similar effect was observed when the interfering stimulus was applied to the ipsilateral little finger, possibly reflecting a process of 'surround inhibition.' Tactile stimulation of more remote regions (principally the face and contralateral hand) resulted in consistent difference wave forms in which the early components (less than 30 msec latency) had scalp distributions differing from one another but consistent with influence on generators in the face or hand region of the second somatosensory cortex (SII). Later potentials consistently identifiable in the difference wave forms were similar for all locations of the interfering stimulus apart from the ipsilateral thumb and were distributed in accordance with a proposed generator in the parietal 'association' cortex.     

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.     

Selective spatial attention induces short-term plasticity in human somatosensory cortex

Early cognitive process in the primary somatosensory cortex (SI) was studied by measuring somatosensory evoked magnetic fields during selective attention tasks. We used vibratory stimuli to the index or middle finger with a frequency of 100 or 400 Hz for selective discrimination of spatial finger and non-spatial frequency attribute of stimuli. An early M50 component from the SI cortex indicated that the SI regions for the fingers were specifically segregated in a finger discrimination task but not in a frequency discrimination task or in a control condition. The task-dependent and immediate switchover of cortical finger representation demonstrates a dynamic SI activation for spatial information processing.     

Functional Organization of the Human First and Second Somatosensory Cortices: a Neuromagnetic Study

Multichannel neuromagnetic recordings were used to differentiate signals from the human first (SI) and second (SII) somatosensory cortices and to define representations of body surface in them. The responses from contralateral SI, peaking at 20 – 40 ms, arose mainly from area 3b, where representations of the leg, hand, fingers, lips and tongue agreed with earlier animal studies and with neurosurgical stimulations and recordings on convexial cortex in man. Representations of the five fingers were limited to a cortical strip of ∼2 cm in length. Responses from SII peaked 100 – 140 ms after contra- and ipsilateral stimuli and varied considerably from one subject to another. Signs of somatotopical organization were seen also in SII. Responses of SII were not fully recovered at interstimulus intervals of 8 s.     

The profile of the recovery cycle in human primary and secondary somatosensory cortex: a magnetoencephalography study.

OBJECTIVE: To estimate the lifetime of sensory memory in human primary (SI) and secondary (SII) somatosensory cortex with a view to furthering our understanding of the roles played by these cortices in the processing of tactile information. METHODS: Somatosensory evoked fields (SEFs) were recorded following trains of 5 electrical pulses applied to the right median nerve at the wrist using a whole-head 80 channel magnetoencephalography (MEG) system. Recordings were acquired for trains of pulses with differing interstimulus intervals (ISIs) occurring at 100, 200, 300, 400 and 500 ms. The profile of SEF intensities for the different ISIs provided an estimate of the recovery cycle of evoked neuronal activity, and the time constant of the exponential curve fitted to the recovery cycle was calculated to obtain a putative measure of the lifetime of somatic sensory memory in SI and SII. RESULTS: The estimated time constants were 0.11+/-0.06 s (mean+/-SD) in SI and 0.82+/-0.34 s in SII. The mean time constant in SII was significantly longer than that in SI (Student's paired t test: P=0.021; analysis of variance: F(1,3)=19.7, P=0.021). CONCLUSIONS: These data indicate that the lifetime of somatic sensory memory is of longer duration in higher order cortical areas than in primary sensory cortex in the somatosensory information processing system.     

Selective attention regulates spatial and intensity information processing in the human primary somatosensory cortex

Attention-related cognitive processes in the primary somatosensory cortex (SI) were studied by measuring somatosensory evoked magnetic fields (SEFs). Twenty-one normal adult human subjects participated in this study for investigating effects of attention and stimulus intensity on cortical finger representation in the SI cortex. Electric stimuli at low and high intensity were delivered to the index or middle finger in finger discrimination and non-discrimination task. For the low intensity stimulation at 1.25 times sensory threshold, an early component (M50) showed clear segregation of the sources for the two fingers and an increase of the amplitude specific to the finger discrimination task. Such an attentional effect on the SI cortex was masked by the high intensity stimulation (2.5 times sensory threshold); the M50 source separation by the fingers was induced irrespective of the discrimination or non-discrimination task. The results suggest that a conscious regulation of stimulus intensity coding in the SI cortex underlies the attention-dependent enhancement of spatial finger information processing.     

Activation of somatosensory cortical areas varies with attentional state: an fMRI study

The differing roles of SI and SII areas in the somatosensory system have received relatively little interest in previous research. In the present study fMRI was applied to determine possible changes in activations of these areas as a function of attentional modulation (attending vs. not attending to the stimulation of a finger). The results showed that attention induced larger regional changes, mostly enlargements of activated areas, at SII than at SI. The number of instances where new, emerging activations, not present in the non-attend condition, were observed was larger at SII than at SI. These differential attentional effects indicate that SII areas may have a role in more complex tactile functions such as tactile working memory mechanisms.     

Sensorimotor integration in human primary and secondary somatosensory cortices

We measured somatosensory evoked fields (SEFs) to electric median nerve stimuli from eight healthy subjects with a whole-scalp 122-channel neuromagnetometer in two different conditions: (i) ‘rest', with stimuli producing clear tactile sensation without any motor movement, and (ii) ‘contraction' with exactly the same stimuli as in ‘rest', but with the subjects maintaining sub-maximal isometric contraction in thenar muscles of the stimulated hand. The aim was to study the role of the primary (SI) and secondary somatosensory (SII) cortices in sensorimotor integration. The amplitude of the SI response N20m did not change with coincident isometric contraction, whereas P35m was significantly reduced. On the contrary, activation of contra- and ipsilateral SII cortices was significantly enhanced during the contraction. We suggest that isometric contraction facilitates activation of SII cortices to tactile stimuli, possibly by decreasing inhibition from the SI cortex. The enhanced SII activation may be related to tuning of SII neurons towards relevant tactile input arising from the region of the body where the muscle activation occurs.     

Magnetic source imaging of tactile input shows task-independent attention effects in SII

We investigated whether attention to different stimulus attributes (location, intensity) has different effects on the activity of the secondary (SII) somatosensory cortex. Tactile stimuli were applied to the left index finger and somatosensory evoked fields (SEFs) were recorded using a whole-head magnetoencephalography (MEG) system. Two oddball paradigms with stimuli varying in location or intensity were performed in an ignore and an attend condition. Brain sources were estimated by magnetic source imaging. No attention effect was observed for the primary SI area. However, attention enhanced SII activity bilaterally from 55 to 130 ms by 52% in the spatial and 64% in the intensity discrimination task. SII attentional enhancement was very similar in both paradigms and occurred both for deviants and standards.     

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.     

Seeing touch in the somatosensory cortex: a TMS study of the visual perception of touch

Recent studies suggest the existence of a visuo-tactile mirror system, comprising the primary (SI) and secondary (SII) somatosensory cortices, which matches observed touch with felt touch. Here, repetitive transcranial magnetic stimulation (rTMS) was used to determine whether SI or SII play a functional role in the visual processing of tactile events. Healthy participants performed a visual discrimination task with tactile stimuli (a finger touching a hand) and a control task (a finger moving without touching). During both tasks, rTMS was applied over either SI or SII, and to the occipital cortex. rTMS over SI selectively reduced subject performance for interpreting whether a contralateral visual tactile stimulus contains a tactile event, whereas SII stimulation impaired visual processing regardless of the tactile component. These findings provide evidence for a multimodal sensory-motor system with mirror properties, where somatic and visual properties of action converge. SI, a cortical area traditionally viewed as modality-specific, is selectively implicated in the visual processing of touch. These results are in line with the existence of a sensory mirror system mediating the embodied simulation concept.     

Altered responses to cutaneous stimuli in the second somatosensory cortex following lesions of the postcentral gyrus in infant and juvenile macaques

Infant macaques recover tactile abilities better than older animals after somatosensory cortical lesions. To investigate the neural basis of this phenomenon, we ablated the hand representation in primary somatosensory cortex (SI) of infant and juvenile Macaca mulatta and recorded in ipsilateral second somatosensory cortex (SII) a year later. We also made tracer injections to verify the lesion boundaries and to study the connections of SII after the lesion of SI. Similar to the report of Pons et al. (Science 237:417-420, '87), we found that substantial portions of the SII hand area were unresponsive to cutaneous stimulation of the hand in both age groups. Particularly, there were no cutaneous receptive fields restricted to the digits. Some responses were elicited in each animal by mechanical stimulation of the hand, including a proportion related to cutaneous receptive fields. This proportion was higher in the infants than in the juveniles, which may explain the greater capacity of the infants for recovery of tactile function after SI lesions. The residual somatic drive in the SII hand area of the juveniles was attributable to sparing of parts of areas 3a and 3b. However, in the infants, this explanation was not tenable since the responses noted in SII occurred even after total ablations of the postcentral gyrus. The pattern of corticocortical connections revealed by injections of HRP into the medial margin of the SI lesion and of Fast Blue into SII in one infant confirmed the absence of SI inputs to the region of SII where responses were recorded from the hand. Representations of body parts other than the hand were normally responsive, and their location was consistent with normal somatotopy in 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.     

Attention modulates beta oscillations during prolonged tactile stimulation

The inter‐play between changes in beta‐band (14–30‐Hz) cortical rhythms and attention during somatosensation informs us about where and when relevant processes occur in the brain. As such, we investigated the effects of attention on somatosensory evoked and induced responses using vibrotactile stimulation and magnetoencephalographic recording. Subjects received trains of vibration at 23 Hz to the right index finger while watching a movie and ignoring the somatosensory stimuli or paying attention to the stimuli to detect a change in the duration of the stimulus. The amplitude of the evoked 23‐Hz steady‐state response in the contralateral primary somatosensory cortex (SI) was enhanced by attention and the underlying dipole source was located 2 mm more medially, indicating top‐down recruitment of additional neuronal populations for the functionally relevant stimulus. Attentional modulation of the somatosensory evoked response indicates facilitation of early processing of the tactile stimulus. Beta‐band activity increased after vibration offset in the contralateral primary motor cortex (MI) [event‐related synchronization (ERS)] and this increase was larger for attended than ignored stimuli. Beta‐band activity decreased in the ipsilateral SI prior to stimulus offset [event‐related desynchronization (ERD)] for attended stimuli only. Whereas attention modulation of the evoked response was confined to the contralateral SI, event‐related changes of beta‐band activity involved contralateral SI–MI and inter‐hemispheric SI–SI connections. Modulation of neural activity in such a large sensorimotor network indicates a role for beta activity in higher‐order processing.