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.     

Spatial properties and interhemispheric differences of the sensory hand cortical representation: a neuromagnetic study

We performed a neuromagnetic investigation of the sensory hand cortical representation in the two hemispheres of 20 healthy volunteers. The localizations within the brain hemispheres of the cortical Equivalent Current Dipoles (ECDs) activated with the shortest latencies (N20 m and P30 m components) by separate stimulation of contralateral median nerve, thumb and little finger were analysed. The ECD spatial coordinates were in agreement with the known somatotopy of the sensory homunculus: little finger more medial and posterior, thumb more lateral and anterior, median nerve in-between. By considering the ECDs to thumb and little finger stimulation the boundaries of the hand cortical representation in primary sensory cortex, the `hand extension' was evaluated as the distance between the two. This parameter was similar on the two hemispheres, the `hand extension' being 17 mm and 12 mm for N20 m and P30 m components, respectively, with a standard deviation of 5 mm. We provide for the first time the ECDs localization of left and right median nerve, thumb and little finger, as well as the `hand extension' values, and their interhemispheric differences as a normative data set describing the organization of primary sensory cortical areas reserved to the hand in the healthy population. This approach permits objective measurements of absolute values, as well as of interhemispheric differences, of the sensory hand area following a monohemispheric lesion as well as to non-invasively follow-up its reorganization during clinical recovery.     

Difference in somatosensory evoked fields elicited by mechanical and electrical stimulations: Elucidation of the human homunculus by a noninvasive method

We recently recorded somatosensory evoked fields (SEFs) elicited by compressing the glabrous skin of the finger and decompressing it by using a photosensor trigger. In that study, the equivalent current dipoles (ECDs) for these evoked fields appeared to be physiologically similar to the ECDs of P30m in median nerve stimulation. We sought to determine the relations of evoked fields elicited by mechanically stimulating the glabrous skin of the great toe and those of electrically produced P40m. We studied SEFs elicited by mechanical and electrical stimulations from the median and tibial nerves. The orientations of dipoles from the mechanical stimulations were from anterior-to-posterior, similar to the orientations of dipoles for P30m. The direction of the dipole around the peak of N20m from median nerve electrical stimulation was opposite to these directions. The orientations of dipoles around the peak of P40m by tibial nerve stimulation were transverse, whereas those by the compression and decompression stimulation of the toe were directed from anterior-to-posterior. The concordance of the orientations in ECDs for evoked fields elicited by mechanical and electrical stimulations suggests that the ECDs of P40m are physiologically similar to those of P30m but not to those of N20m. The discrepancy in orientations in ECDs for evoked field elicited by these stimulations in the lower extremity suggests that electrical and compression stimulations elicit evoked fields responding to fast surface rubbing stimuli and/or stimuli to the muscle and joint.     

Neural connectivity in hand sensorimotor brain areas: an evaluation by evoked field morphology

The connectivity pattern of the neural network devoted to sensory processing depends on the timing of relay recruitment from receptors to cortical areas. The aim of the present work was to uncover and quantify the way the cortical relay recruitment is reflected in the shape of the brain-evoked responses. We recorded the magnetic somatosensory evoked fields (SEF) generated in 36 volunteers by separate bilateral electrical stimulation of median nerve, thumb, and little fingers. After defining an index that quantifies the shape similarity of two SEF traces, we studied the morphologic characteristics of the recorded SEFs within the 20-ms time window that followed the impulse arrival at the primary sensory cortex. Based on our similarity criterion, the shape of the SEFs obtained stimulating the median nerve was observed to be more similar to the one obtained from the thumb (same median nerve innervation) than to the one obtained from the little finger (ulnar nerve innervation). In addition, SEF shapes associated with different brain regions were more similar within an individual than between subjects. Because the SEF morphologic characteristics turned out to be quite diverse among subjects, we defined similarity levels that allowed us to identify three main classes of SEF shapes in normalcy. We show evidence that the morphology of the evoked response describes the anatomo-functional connectivity pattern in the primary sensory areas. Our findings suggest the possible existence of a thalamo-cortico-thalamic responsiveness loop related to the different classes.     

Somatosensory evoked magnetic fields from the primary somatosensory cortex (SI) in acute stroke.

We recorded somatosensory evoked magnetic fields (SEFs) to median nerve stimulation from 15 patients in the acute stage (1-15 days from the onset of the symptoms) of their first-ever unilateral stroke involving sensorimotor cortical and/or subcortical structures in the territory of the middle cerebral artery (MCA). Neuronal activity corresponding to the peaks of the N20m, P35m and P60m SEF deflections from the contralateral primary somatosensory cortex (SI) was modelled with equivalent current dipoles (ECDs), the locations and strengths of which were compared with those of an age-matched normal population. Four patients with pure motor stroke had symmetric SEFs. In one of the 4 patients with pure sensory stroke, and in 5 of the 7 patients with sensorimotor paresis, the SEFs were markedly attenuated or missing. All except one patient with abnormal SEFs had deficient two-point discrimination ability; especially the attenuation of N20m was more clearly correlated with two-point discrimination than with joint-position or vibration senses. Of the different SEF deflections, P35m and P60m were slightly more sensitive indicators of abnormality than N20m, the former being affected in two patients with symmetric N20m. Three patients with pure sensory stroke and lesions in the opercular cortex had normal SEFs from SI. We conclude that the SEF deflections N20m, P35m and P60m from SI are related to cutaneous sensation, in particular discriminative to touch. The results also demonstrate that basic somatosensory perception can be affected by lesions in the opercular cortex in patients with functionally intact SI.     

Magnetoencephalographic investigation of somatosensory homunculus in patients with peri-Rolandic tumors

In order to investigate functional topography of the hand somatosensory cortex in five patients with peri-Rolandic tumors (four frontal lobes and one parietal lobe), we recorded somatosensory evoked fields (SEFs) using magnetoencephalography (MEG) after stimulation of the median nerve (MN) and the five digits. The results obtained were compared with those of five normal healthy subjects. In all five patients, SEFs following MN and digit stimulation showed the previously described respective N20m and N22m components of primary sensory response. Single dipole modeling was applied to determine the three dimensional cortical representations of the N20m and N22m components. The cortical representations of the hand were identical to those of normal subjects, arranging in an orderly somatotopic way from lateral inferior to medial superior in the sequence thumb, MN, index, middle, ring, and little fingers. This sensory homunculus was confirmed by cortical recording of the somatosensory evoked potentials (SEPs) at the time of surgery. Thus, we demonstrate that SEFs, recorded on MEG in conjunction with source localization techniques, are useful to non-invasively investigate the functional topography of the human hand somatosensory cortex in pathological conditions.     

The temporal profile of interactions between sensory information from both hands in the secondary somatosensory cortex.

OBJECTIVE: To elucidate the temporal profile of interactions between sensory information from both hands in the somatosensory cortex. METHODS: Somatosensory evoked fields (SEFs), generated by stimulation applied to the right index finger after a preceding stimulation to the left index finger, were recorded using a whole head-type magnetoencephalography (MEG). The paired electrical stimuli were applied with a stimulation onset asynchrony (SOA) of 50, 100, 200, 300, or 400 ms. RESULTS: The mean SEF intensities in the primary somatosensory area (SI) of five subjects, which were evoked approximately 40 ms after the latter of the paired stimuli, were not significantly smaller than that evoked in the control condition when only the right finger was stimulated. In contrast, SEFs in the secondary somatosensory area (SII), generated approximately 100 ms after the stimuli, were suppressed when the paired stimuli were applied at an SOA of 100 ms (P<0.05, t test). In addition, SEFs at approximately 150 ms after the stimuli were significantly suppressed at SOAs of 50, 100 (P<0.05), 200, and 300 ms (P<0.1). CONCLUSION: Within a time window of approximately 300 ms, sensory information from the left finger significantly affected the SEFs generated by sensory inputs from the right finger. This time window may be required for the integration of sensory input.     

The effect of tourniquet-induced ischemia on somatically evoked cerebral magnetic fields in man

We recorded somatically evoked magnetic fields (SEFs) during tourniquet-induced ischemia in healthy humans. The subjective intensities of the stimuli decreased earlier and more clearly than the amplitudes of the SEF deflections peaking at 60-80 ms. Mixed nerve and sensory nerve responses behaved differently during ischemia. The results indicate that these SEF deflections are transmitted through thick myelinated fibers. Muscle afferents might contribute considerably to mixed nerve SEFs.     

Diagnostic specificity of sensory and motor nerve conduction variables in early detection of carpal tunnel syndrome

Summary In the carpal tunnel syndrome (CTS) sensory nerve conduction is more sensitive than motor conduction. However, 8%–25% of the sensory distal latencies in symptomatic hands may still be normal. A systematic study was made of the median, ulnar and radial orthodromic nerve conduction velocities (SNCV) stimulating each of the fingers separately. Four SNCVs from the median nerve, two SNCVs from the ulnar nerve and one from the radial nerve were obtained, and the ratio of the median to radial SNCV and the ratios of the median and ulnar SNCVs were estimated. The significance of these parameters in the diagnosis of the CTS was studied, and a rapid technique for the screening of nerve entrapment in the initial stages of the disease is proposed. Three hundred and seventy-five symptomatic hands were examined. Seventy-five hands showed normal distal latency, in which cases, however, the SNCV of the ring finger was always outside the normal range, while the SNCVs of the thumb, index and middle fingers were abnormal in 64%, 80% and 92% of cases respectively. The amplitudes of the sensory responses were the least sensitive of the parameters studied. Our results suggest that a study of the median nerve digital branch to the ring finger may be of value in providing an easily performed and rapid technique for screening an early median nerve entrapment at the wrist.     

Differential interaction of somatosensory inputs in the human primary sensory cortex: a magnetoencephalographic study.

OBJECTIVE: Somatosensory evoked magnetic fields (SEFs) were recorded to investigate the interaction of the somatosensory inputs using the modality of electrical finger stimulation in 6 normal subjects. METHODS: Electrical stimuli were given to the index (II), middle (III) or little (V) fingers individually, and also to pairs of either the II and III simultaneously, or the II and V simultaneously. The interaction ratio (IR) was calculated as the ratio of the SEF amplitude by simultaneous two-finger stimulation to the arithmetically summed SEF amplitudes of two individual-finger stimulations. RESULTS: SEFs showed 3 major components: N22m, P30m and P60m. The N22m and P60m revealed a clear somatotopic organization in the primary sensory cortex (S1) in the sequence of II, III and V, while the P30m showed a cluster with medial location compared with N22m and P60m in S1. The N22m had a significantly greater IR in II and III stimulation compared to that in II and V stimulation. The P60m also showed a similar trend in the IR but was greater than that of N22m. In contrast, the IR in P30m showed no such tendency. CONCLUSION: The interaction of S1 was most influenced when adjacent receptive fields were activated in the modality of electrical finger stimulation. Our results were consistent with the concept that the Brodmann's areas in S1 which produce the 3 components of the SEFs have different functional organization.     

Rapid functional plasticity in the primary somatomotor cortex and perceptual changes after nerve block

The mature human primary somatosensory cortex displays a striking plastic capacity to reorganize itself in response to changes in sensory input. Following the elimination of afferent return, produced by either amputation, deafferentation by dorsal rhizotomy, or nerve block, there is a well-known but little-understood 'invasion' of the deafferented region of the brain by the cortical representation zones of still-intact portions of the brain adjacent to it. We report here that within an hour of abolishing sensation from the radial and medial three-quarters of the hand by pharmacological blockade of the radial and median nerves, magnetic source imaging showed that the cortical representation of the little finger and the skin beneath the lower lip, whose intact cortical representation zones are adjacent to the deafferented region, had moved closer together, presumably because of their expansion across the deafferented area. A paired-pulse transcranial magnetic stimulation procedure revealed a motor cortex disinhibition for two muscles supplied by the unaffected ulnar nerve. In addition, two notable perceptual changes were observed: increased two-point discrimination ability near the lip and mislocalization of touch of the intact ulnar portion of the fourth finger to the neighbouring third finger whose nerve supply was blocked. We suggest that disinhibition within the somatosensory system as a functional correlate for the known enlargement of cortical representation zones might account for not only the 'invasion' phenomenon, but also for the observed behavioural correlates of the nerve block.     

拇指感觉神经传导速度诊断轻度腕管综合征的临床应用价值

目的:探讨拇指感觉神经传导速度(sensory nerve conduction velocity,SCV)诊断轻度腕管综合征的临床应用价值。方法:对18例(26只手)轻度腕管综合征的患者和15例(30只手)年龄性别相匹配的正常人,测定了腕部正中神经和桡神经的感觉神经传导速度,并进行对比研究。结果:中指正中神经SCV的异常率为50%,腕部正中神经/桡神经(刺激拇指)SCV差值的异常率为84.6%,明显大于用常规检查方法组(刺激中指),有8例12只手腕部正中神经感觉动作电位(刺激拇指)中出现双峰电位,而对照组则无。结论:在腕管综合征肌电图的诊断中,比较正中神经和桡神经SCV的差值是早期诊断腕管综合征的敏感指标之一。     

Cortical sensory representation of the human hand: size of finger regions and nonoverlapping digit somatotopy.

Findings differ on cortical representation of fingers between human and animal studies, and on digit somatotopy among human studies. To resolve these differences, we mapped cortical sensory representation of each of the five digits and of median and ulnar nerves in three patients, using focal peripheral electrical shock stimuli. We compared locations and sizes of cortical regions among digits and nerves, using the model of a current dipole in a sphere applied to electrocorticography from subdural grids. Cortical representation was larger for the index finger than for the little finger and for the middle finger than for the ring finger, which are similar to findings in the monkey but different from Penfield's classic sensory homunculus. The thumb was larger than the middle finger, as in the homunculus. There was nonoverlapping somatotopy of all digits in each patient. These findings demonstrate a previously unrecognized similarity of cortical sensory organization of the fingers between humans and other primates.     

Orthodromic sensory conduction along the ring finger in normal subjects and in patients with a carpal tunnel syndrome.

The purpose of the present study was to examine the value of measuring sensory conduction along the median and ulnar nerves of the fourth finger in the diagnosis of a carpal tunnel syndrome (CTS). In 23 controls, sensory conductions along median and ulnar nerves were identical. In 28 of 38 patients with CTS, stimulation of the ring finger revealed a reduced conduction velocity along sensory median nerve fibres in contrast to normal conduction along ulnar sensory nerve fibres. In 5 patients, a sensory action potential was absent over the median nerve and in another 5 sensory conduction was normal along both nerves. We conclude that testing of sensory conduction along the ring finger is useful in about 74% of patients with CTS, while in the remaining 26% other fingers must be examined to establish the diagnosis.     

Evaluation of the dynamics of sensory improvement in the hand after surgical treatment of carpal tunnel syndrome

BACKGROUND AND PURPOSE: The aim of the study was to evaluate the dynamics of improvement of tactile sensory threshold and two-point discrimination in fingers innervated by the median nerve after surgical treatment of carpal tunnel syndrome (CTS), to assess differences between improvements of both aforementioned sensory measures, as well as to estimate the time taken to achieve ultimate sensory improvement in the hand. MATERIAL AND METHODS: The study included 87 patients (78 females and 9 males) operated on at the Department of Traumatology and Hand Surgery, Medical Academy in Wroclaw between July 2002 and August 2004 because of carpal tunnel syndrome. Mean age of patients was 51.8 years. The study included evaluation of tactile sensory (pressure) threshold and two-point discrimination. Measurements were performed seven times in each patient, that is, before the surgery, and then on the second, fifth and tenth postoperative day as well as one, three and six months after the surgery. RESULTS: There were only minor differences in improvement of tactile sensory threshold and two-point discrimination between particular fingers, but in each case less favourable results were achieved in the thumb and the middle finger and the best results in the ring finger. Over time the observed degree of sensory improvement decreased, and during the last follow-up mean values of tactile sensory threshold and two-point discrimination in each of the studied fingers were within the normal range. CONCLUSIONS: After surgical treatment of CTS normalisation of two-point discrimination occurs earlier than normalisation of tactile sensory threshold. The highest degree of improvement of two-point discrimination is observed during the first two days after surgery, while the highest degree of improvement of tactile sensory threshold is noted between the 10th and 30th postoperative day. No further significant improvement of tactile sensory threshold or two-point discrimination is observed following an initial period of three months after surgery.     

Altered Finger Representations in Sensorimotor Cortex of Musicians with Focal Dystonia: Precentral Cortex

Using functional magnetic resonance imaging (fMRI), finger representations were characterized in the precentral cortex of 11 normal musicians and 14 musicians with focal task-specific dystonia. Finger representations were identified from differential activation during repetitive movements of each finger relative to others. Despite group similarities in topography, abnormalities in representations of affected fingers were identified. For the finger showing chronic flexion (primary dystonic finger or PDF), the cortical “disparity” from its normal location and the distance to the adjacent finger were increased. By contrast, representational characteristics of the finger showing chronic extension (primary compensatory finger or PCF) did not differ significantly from the control group, but did differ from those of the PDF. Regardless of whether either finger's representation differed substantively from normal, the PCF consistently showed greater volume of activation than the PDF or other fingers. These findings reflect dysfunctional interactions between at least two fingers and their cortical representations.     

Maturation of somatosensory cortical processing from birth to adulthood revealed by magnetoencephalography

ObjectiveTo evaluate the maturation of tactile processing by recording somatosensory evoked magnetic fields (SEFs) from healthy human subjects.MethodsSEFs to tactile stimulation of the left index finger were measured from the contralateral somatosensory cortex with magnetoencephalography (MEG) in five age groups: newborns, 6- and 12–18-month-olds, 1.6–6-year-olds, and adults. The waveforms of the measured signals and equivalent current dipoles (ECDs) were analyzed in awake and sleep states in order to separate the effects of age and vigilance state on SEFs.ResultsThere was an orderly, systematic change in the measured and ECD source waveforms of the initial cortical responses with age. The broad U-shaped response in newborns (M60) shifted to a W-shaped response with emergence of a notch by 6 months of age. The adult-type response with M30 and M50 components was present by 2 years. The ECDs of M60 and M30 were oriented anteriorly and that of M50 posteriorly. These maturational changes were independent of vigilance state.ConclusionsThe most significant maturation of short latency cortical responses to tactile stimulation takes place during the first 2 years of life.SignificanceThe maturational changes of somatosensory processing can noninvasively be evaluated with MEG already in infancy.     

Two evoked responses with different recovery functions in the primary somatosensory cortex in humans.

OBJECTIVES: We investigated the recovery function of somatosensory evoked magnetic cortical fields (SEFs) to confirm the temporal aspects of the somatosensory process in humans. METHODS: SEFs were recorded following median nerve electrical stimulation in 6 healthy subjects. Double stimulation, with interstimulus intervals (ISIs) from 3 to 100 ms, was applied, and the SEF components for the second stimulation were analyzed. In a supplementary experiment, responses to single stimulations of various intensities from the sensory threshold to the motor threshold were studied. RESULTS: The first SEF component (1M) diminished when the ISI was less than 10 ms, while the second component (2M) remained even when the ISI was 3 ms. The two components showed a very similar attenuation with decrease of stimulus intensity. There was no significant difference in dipole location between 1M and 2M in the primary somatosensory cortex (SI). CONCLUSIONS: The results suggested that at least two independent pathways with different recovery functions exist in a similar area in the SI.     

Specific somatosensory processing in somatosensory area 3b for human thumb: a neuromagnetic study.

OBJECTIVES: We examined the relation between somatosensory N20m primary responses and high-frequency oscillations (HFOs) after thumb and middle finger stimulation. METHODS: Somatosensory evoked fields (SEFs) from 12 subjects were measured following electric stimulation of the thumb and middle finger. SEFs were recorded with a wide bandpass (3-2000 Hz) and then N20m and HFOs were separated by subsequent 3-300 and 300-900 Hz bandpass filtering. RESULTS: The N20m peak-to-peak amplitude did not differ significantly between thumb and middle finger SEFs. In contrast, HFOs had a significantly larger number of peaks and were higher in the maximum amplitude and the total amplitude after thumb stimulation than after middle finger stimulation. CONCLUSIONS: Our present data demonstrate a different relation between N20m and HFOs after thumb and middle finger stimulation. In view of the fact that the human thumb has uniquely evolved functionally and morphologically, the somatosensory information from the thumb will be processed differently for a fine motor control. We speculate that HFOs are generated by inhibitory interneurons in layer 4 in area 3b. Thus, enhanced activity of interneurons reflected by high amplitude HFOs exerts stronger inhibition on downstream pyramidal cells in area 3b for thumb stimulation.     

District-related frequency specificity in hand cortical representation: dynamics of regional activation and intra-regional synchronization

The aim of this work was to study the degree of neuronal synchronization occurring within the portion of the somatosensory cortex devoted to hand control during an external sensory stimulation. In this way, we focused on the properties of the sensory cortical representation, rather than the more investigated motor one. To this aim, we collected magnetoencephalograhic data from healthy subjects during separate stimulation of their thumbs and little fingers and analyzed these data by means of a time-dependent 'synchronization index'. The properties of this index within the beta [16-32 Hz] and gamma [36-44 Hz] frequency bands suggest that the hand representation in the human primary cortex follows a frequency coding, in addition to the somatotopic one, for discriminating different districts. Our results showed that the gamma synchronization is higher following stimulation of the thumb than of the little finger and we suggest that the strength of gamma band synchronization works as a code for functional prevalence. In particular, our comparative analysis of the dynamic synchronization index and the signal amplitude suggests that a prevalent district (thumb) recruits a smaller number of higher-synchronic gamma band tuned neurons than a non-prevalent district (little finger).