Spatial attention modulates early face processing

It is widely reported that inverting a face dramatically affects its recognition. Previous studies have shown that face inversion increases the amplitude and delays the latency of the face-specific N170 component of the event-related potential (ERP) and also enhances the amplitude of the occipital P1 component (latency 100–132 ms). The present study investigates whether these effects of face inversion can be modulated by visual spatial attention. Participants viewed two streams of visual stimuli, one to the left and one to the right of fixation. One stream consisted of a sequence of alphanumeric characters at 6.67 Hz, and the other stream consisted of a series of upright and inverted images of faces and houses presented in randomized order. The participants’ task was to attend selectively to one or the other of the streams (during different blocks) in order to detect infrequent target stimuli. ERPs elicited by inverted faces showed larger P1 amplitudes compared to upright faces, but only when the faces were attended. In contrast, the N170 amplitude was larger to inverted than to upright faces only when the faces were not attended. The N170 peak latency was delayed to inverted faces regardless of attention condition. These inversion effects were face specific, as similar effects were absent for houses. These results suggest that early stages of face-specific processing can be enhanced by attention, but when faces are not attended the onset of face-specific processing is delayed until the latency range of the N170.     

Electrical somatosensory stimulation modulates hand motor function in healthy humans

In healthy people, electrical somatosensory stimulation modulates excitability in contralateral cortical motor areas. The question whether this is associated with a change in motor performance is still under debate. The effect of electrical somatosensory stimulation on motor performance of the hand was investigated in 14 healthy right-handed subjects. Subjects performed index finger and hand tapping movements as well as reach-to-grasp movements towards small and large cubes with each hand prior to (baseline condition) and following 2-hour electrical somatosensory stimulation (trains of 5 pulses at 10 Hz with 1 ms duration delivered at 1 Hz with an intensity on average 60 % above the individual somatosensory threshold) of the (i) right median nerve, (ii) left median nerve, (iii) right tibial nerve (control stimulation) and (iv) left tibial nerve (control stimulation) on separate occasions at least one week apart. The order of sessions was counterbalanced across subjects. Somatosensory stimulation of the median nerves, but not of the tibial nerves, reduced the frequency and velocity of index finger and hand tapping movements performed with the stimulated hand, compared to baseline. In contrast, the kinematics of reach-to-grasp movements remained unaffected by somatosensory stimulation. The data suggest that somatosensory stimulation interferes with the processing of highly automated open-loop motor output at the stimulated limb, as reflected by tapping movements, but not with the processing of closed-loop motor performance, as reflected by reach-to-grasp movements.     

Spatial attention determines the nature of nonverbal number representation

Coordinated studies of adults, infants, and nonhuman animals provide evidence for two systems of nonverbal number representation: a "parallel individuation" system that represents individual items and a "numerical magnitude" system that represents the approximate cardinal value of a group. However, there is considerable debate about the nature and functions of these systems, due largely to the fact that some studies show a dissociation between small (1-3) and large (>3) number representation, whereas others do not. Using event-related potentials, we show that it is possible to determine which system will represent the numerical value of a small number set (1-3 items) by manipulating spatial attention. Specifically, when attention can select individual objects, an early brain response (N1) scales with the cardinal value of the display, the signature of parallel individuation. In contrast, when attention cannot select individual objects or is occupied by another task, a later brain response (P2p) scales with ratio, the signature of the approximate numerical magnitude system. These results provide neural evidence that small numbers can be represented as approximate numerical magnitudes. Further, they empirically demonstrate the importance of early attentional processes to number representation by showing that the way in which attention disperses across a scene determines which numerical system will deploy in a given context.     

Hand posture modulates neuronal interaction in the primary somatosensory cortex of humans.

OBJECTIVE: To investigate the effects of hand posture on the modulation of neuronal interactions in the cortical finger regions of the human somatosensory cortex. METHODS: Neuronal magnetic fields, evoked by electrical stimuli to the thumb and/or to the index finger of the right hand, were recorded in different hand postures ('OPEN': opened hand and 'CLOSE': both fingers in opposite position to pick up something) by using a whole head type magnetoencephalography. The equivalent current dipole (ECD) for components in the primary (SI) and secondary somatosensory cortices (SII) was calculated. The interaction ratio (IR) was calculated as a ratio of the vector sum of ECD moments evoked by respective stimulation of each finger to the ECD moment evoked by simultaneous stimulation of both fingers. RESULTS: The mean IR of N20m was significantly larger in CLOSE than in OPEN (p=0.033, ANOVA). On the contrary, the IR of P40m was larger in OPEN than in CLOSE (p=0.042). The IR of SII components was not significantly different between the different hand postures (p=0.35). CONCLUSIONS: Neuronal interaction between the thumb and index finger in the human SI is modulated by hand posture. Provided that forming hand posture is related to receiving sensory input, the interaction modulation may play a role in the facilitation of somatosensory processing. SIGNIFICANCE: Our results suggest experimental evidence for the immediate modulation of neuronal activity in the somatosensory area.     

Myelin stains reveal an anatomical framework for the representation of the digits in somatosensory area 3b of macaque monkeys

Brain sections cut parallel to the cortical surface revealed myelin-light septa that isolated representations of the digits and parts of the face, teeth, and tongue in area 3b of adult and infant macaque monkeys. The widths of the bands of cortex representing individual digits, as measured by the distances between isolating septa, were proportionally similar in infant (2-4 week) and adult monkeys. However, the bands for digits 1-3 were somewhat narrower in infant than adult monkeys. There was little variation in absolute widths across individuals in the infant or adult groups, or between left and right hemispheres of the same group. Widths for digits 1-4 progressively decreased. The results suggest that these isomorphs of digits emerge in prenatal or early postnatal development and typical variations in postnatal hand use have little impact on subsequent development. As the hand representation in somatosensory cortex of monkeys may be significantly altered after the partial loss of peripheral nerve inputs, the physiological representation is not completely constrained by the isolating septa. Instead, the septa may serve as a persistent marker of normal organization in studies of cortical reorganization.     

Somatotopic representation of pain in the primary somatosensory cortex (S1) in humans

ObjectiveIn contrast to tactile inputs, the organization and processing of nociceptive inputs in the primary somatosensory cortex (S1) remain largely unexplored. Few studies have examined the arrangement of nociceptive inputs in S1. The aim of this study was to investigate the representation of nociceptive inputs in the human cortex, including the somatosensory and posterior parietal cortices, from widely separated cutaneous sites.MethodsWe examined the somatotopic organization of the nociceptive system in S1, opercular and posterior parietal cortices by measuring the magnetoencephalographic responses (somatosensory-evoked magnetic fields) of four healthy controls in response to intraepidermal electrical stimulation applied to the face, neck, back, elbow, wrist, hand, finger, knee, and foot, which selectively activated the Aδ fibers.ResultsMagnetoencephalography demonstrated clear somatotopy in the S1 responses to noxious stimuli, with the foot representation in the extreme posteromedial position of S1 and the facial area in the extreme anterolateral position. There was little evidence of any clear somatotopic organization in the secondary somatosensory and posterior parietal cortices.ConclusionThese findings suggest that the nociceptive system uses the large body surface map in S1.SignificanceThis is the first MEG study to demonstrate the cortical representation of nociceptive inputs in the human S1. We showed that widely separated cutaneous sites clearly supported Penfield’s homunculus.     

Spatial coincidence modulates interaction between visual and somatosensory evoked potentials

The time course of interaction between concurrently applied visual and somatosensory stimulation with respect to evoked potentials (EPs) was studied. Visual stimuli, either in the left or right hemifield, and electric stimuli to the left wrist were delivered either alone or simultaneously. Visual and somatosensory EPs were summed and compared to bimodal EPs (BiEP, response to actual combination of both modalities). Temporal coincidence of stimuli lead to sub-additive or over-additive amplitudes in BiEPs in several time windows between 75 and 275 ms. Additional effects of spatial coincidence (left wrist with left hemifield) were found between 75 and 300 ms and beyond 450 ms. These interaction effects hint at a temporo-spatial pattern of multiple brain areas participating in the process of multimodal integration.     

Multiplicity in the high-frequency signals during the short-latency somatosensory evoked cortical activity in humans.

OBJECTIVE: Recent studies using electroencephalography or magnetoencephalography have shown that peripheral nerve stimulations produce short-latency high-frequency signals in the human somatosensory cortex. The present study tested whether they consist of more than one distinct type of signal. METHODS: Somatic evoked magnetic fields (SEFs) elicited by electrical stimulation of the median nerve were measured in 12 healthy volunteers. They were analyzed using a time-frequency analysis method based on Gabor filters and another based on autoregressive moving average, and also with bispectrum and bicoherence techniques and a new dispersion curve method. RESULTS: Signals in two separate high-frequency bands (200 and 600 Hz) were distinguished from the main signal in the low frequency (LF) range during the time period of N20m and P25m. The novel 200 Hz-band signal was seen reliably in those channels where the LF band signal was weak, so that the former was not masked by the latter. The 600 Hz signal consisted of two distinct components or parts (p1 and p2) in 10 out of 12 subjects, one peaking during ascending slope and the second during the descending slope of the N20m. The latency of the p1 was shorter than the latencies of the 200 Hz and LF signals according to the dispersion curve analysis. The inter-peak interval of p1 became shorter for later peaks in all 12 subjects. Bicoherence analysis revealed a significant phase coupling between the 200 and 600 Hz bands. CONCLUSIONS: There are three distinct types of signal during the time period of the short-latency cortical components of the SEF -- LF which gives rise to the commonly seen waveform of the SEF, the newly found 200 Hz signal and the 600 Hz signal which consists of two components. The possible origins of the high frequency signals are discussed in light of the new set of evidence found in the present study.     

Predictability of the target stimulus for sensory-guided movement modulates early somatosensory cortical potentials.

OBJECTIVE: To investigate the role of sensory modulation in the control of sensory-guided behaviour. Specifically, we hypothesized that early somatosensory evoked potentials (SEPs) would be facilitated during performance of continuous sensory-guided movement requiring sustained attention. METHODS: Median nerve SEPs were elicited via electrical stimulation and recorded from scalp electrodes while subjects performed tasks requiring continuous sensory-motor transformations. Subjects received a predictable (rhythmic amplitude modulation) or unpredictable (random amplitude modulation) amplitude varying tactile stimulus (frequency constant at 20 Hz) delivered to the tip of the index finger either alone or with the requirement to track it by modulating the isometric grip force produced by the opposite hand. RESULTS: Early SEP (N20-P27) amplitudes were differentially modulated during unpredictable tracking compared to sensory-motor controls. Specifically, N20 amplitudes were attenuated and P27 amplitudes were enhanced during sensory-guided tracking. CONCLUSIONS: Sustained attention to task-relevant sensory stimuli differentially modulates areas within primary somatosensory cortex (S1) during a continuous sensory-motor transformation. SIGNIFICANCE: These data have implications for understanding the role of attention in regulating somatosensory cortices during sensory-motor behaviour.     

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.     

Selective spatial attention to left or right hand flutter sensation modulates the steady-state somatosensory evoked potential

Steady-state somatosensory evoked potentials (SSSEPs) were recorded from the scalp of human subjects elicited by 20 and 26 Hz mechanical vibrations applied simultaneously to the index finger of the left (20 Hz) and right hand (26 Hz). Subjects were instructed to attend to the flutter vibration at one finger while ignoring the other finger and to detect rare target events at the to-be-attended finger. The amplitude of the frequency coded SSSEP elicited by the attended vibration was significantly enlarged when attention was focused at the respective finger. This amplitude enhancement with attention was most prominent over fronto-central electrode locations contralateral to the attended finger. This is the first report to show the attentional modulation of the SSSEP amplitude in humans, suggesting an enhancement of neural responses in the sense of flutter with attention. The findings will open a new approach for studying the neural mechanisms of sustained selective attention in somatosensation.     

Spatial representation and attention in toddlers with Williams syndrome and Down syndrome

The nature of the spatial representations that underlie simple visually guided actions early in life was investigated in toddlers with Williams syndrome (WS), Down syndrome (DS), and healthy chronological age- and mental age-matched controls, through the use of a "double-step" saccade paradigm. The experiment tested the hypothesis that, compared to typically developing infants and toddlers, and toddlers with DS, those with WS display a deficit in using spatial representations to guide actions. Levels of sustained attention were also measured within these groups, to establish whether differences in levels of engagement influenced performance on the double-step saccade task. The results showed that toddlers with WS were unable to combine extra-retinal information with retinal information to the same extent as the other groups, and displayed evidence of other deficits in saccade planning, suggesting a greater reliance on sub-cortical mechanisms than the other populations. Results also indicated that their exploration of the visual environment is less developed. The sustained attention task revealed shorter and fewer periods of sustained attention in toddlers with DS, but not those with WS, suggesting that WS performance on the double-step saccade task is not explained by poorer engagement. The findings are also discussed in relation to a possible attention disengagement deficit in WS toddlers. Our study highlights the importance of studying genetic disorders early in development.     

Inhibition of cortical somatosensory processing during and after low frequency peripheral nerve stimulation in humans

ObjectiveTranscutaneous low-frequency stimulation (LFS) elicits long-term depression-like effects on human pain perception. However, the neural mechanisms underlying LFS are poorly understood. We investigated cortical activation changes occurring during LFS and if changes were associated with reduced nociceptive processing and increased amplitude of spontaneous cortical oscillations post-treatment.MethodsLFS was applied to the radial nerve of 25 healthy volunteers over two sessions using active (1 Hz) or sham (0.02 Hz) frequencies. Changes in resting electroencephalography (EEG) and laser-evoked potentials (LEPs) were investigated before and after LFS. Somatosensory-evoked potentials were recorded during LFS and source analysis was carried out.ResultsIpsilateral midcingulate and operculo-insular cortex source activity declined linearly during LFS. Active LFS was associated with attenuated long-latency LEP amplitude in ipsilateral frontocentral electrodes and increased resting alpha (8–12 Hz) and beta (16–24 Hz) band power in electrodes overlying operculo-insular, sensorimotor and frontal cortical regions. Reduced ipsilateral operculo-insular cortex source activity during LFS correlated with a smaller post-treatment alpha-band power increase.ConclusionsLFS attenuated somatosensory processing both during and after stimulation.SignificanceResults further our understanding of the attenuation of somatosensory processing both during and after LFS.     

Spatial attention modulates activity in a posterior "where" auditory pathway

Selectively listening to a single location in space modulates both the behavioral and electrophysiological responses to auditory stimuli presented at that location. Transient attention oriented in cue-target or target-target paradigms results in several modulations of the auditory event-related potential known as the Nd1, Nd2, and Nd3. By employing electrical source analysis we tested the hypothesis that the earliest component (the Nd1) reflects modulation of neurons in parietal rather than auditory cortex. It was found that the most likely sources of the Nd1 modulation were posterior to primary auditory cortex within or near the temporo-parietal junction (TPJ). This location is within the putative auditory "where" pathway.     

Physiological changes in the somatosensory forepaw cerebral cortex of adult raccoons following lesions of a single cortical digit representation

The purpose of this study was to determine whether restricted lesions within primary somatosensory (SmI) cortex cause changes in the functional organization of cortical areas bordering on the site of injury. Focal ablations of cortical tissue were made in the representational area for digit 3 within the SmI forepaw cortex of adult raccoons. Electrophysiological mapping experiments done 15-17 weeks later showed that significant alterations had occurred in the response properties of clusters of neurons within those representational zones adjoining the lesion--the zones for digit 2, digit 4, and the palmar pads. These three cortical areas were modified by the appearance of new, usually weaker secondary inputs and changes in some properties of the normal primary inputs from the forepaw. (i) Many neurons responded to stimulation of previously ineffective skin regions; the new inputs often originated from digit 3 but frequently involved other digits or the pads as well. (ii) Neuronal receptive fields (RFs), mapped at a standard suprathreshold stimulus intensity, were larger than normal. (iii) Skin type and submodality sensitivity typically were less specific than normal; more neurons had RFs that included both glabrous and hairy skin or claws and displayed mixtures of responsiveness to skin touch, hair deflection, or claw touch. (iv) The representation of RF location, skin type, and submodality sensitivity was more variable as a function of horizontal and vertical distance through the cortex. In general, the physiological changes were found to degrade the somatotopic order and response specificity of the intact cortical areas adjoining the lesion.(ABSTRACT TRUNCATED AT 250 WORDS)     

Seeing touch and pain in a stranger modulates the cortical responses elicited by somatosensory but not auditory stimulation

Viewing other's pain inhibits the excitability of the motor cortex and also modulates the neural activity elicited by a concomitantly delivered nociceptive somatosensory stimulus. As the neural activity elicited by a transient nociceptive stimulus largely reflects non nociceptive‐specific, multimodal neural processes, here we tested, for the first time, whether the observation of other's pain preferentially affects the brain responses elicited by nociceptive stimulation, or instead similarly modulates those elicited by stimuli belonging to a different sensory modality. Using 58‐channel electroencephalography (EEG), we recorded the cortical responses elicited by laser and auditory stimulation during the observation of videoclips showing either noxious or non‐noxious stimulation of a stranger's hand. We found that the observation of other's pain modulated the cortical activity consisting in an event‐related desynchronization in the β band (β ERD), and elicited by nociceptive laser stimuli, but not by auditory stimuli. Using three different source analysis approaches, we provide converging evidence that such modulation affected neural activity in the contralateral primary sensorimotor cortex. The magnitude of this modulation correlated well with a subjective measure of similarity between the model's hand and the onlooker's representation of the hand. Altogether, these findings demonstrate that the observation of other's pain modulates, in a somatosensory‐specific fashion, the cortical responses elicited by nociceptive stimuli in the sensorimotor cortex contralateral to the stimulated hand. Hum Brain Mapp, 2012. © 2012 Wiley Periodicals, Inc.     

Anomalous somatosensory cortical localization in schizophrenia

OBJECTIVE: Schizophrenia has been reported to be associated with altered localization of regions responding to sound in the auditory cortex, suggesting possible cortical reorganization accompanying the disorder, but it is not clear whether such disturbances are present in any primary sensory cortex other than the auditory. This study examines magnetoencephalographic localization of field sources evoked by tactile stimulation generated in area 3b of the primary somatosensory cortex in patients with schizophrenia compared with normal subjects. METHOD: A total of 29 subjects participated; 14 were patients with schizophrenia, and 15 were normal comparison subjects. Equivalent current dipole generators in both hemispheres were used to model the sources of the 50-msec latency somatosensory evoked field (M50) components produced by contralateral tactile mechanoreceptor stimulation of the tip of the index finger. RESULTS: Compared with normal subjects, patients with schizophrenia demonstrated reversed asymmetry (left anterior to right) and relatively anterior and inferior displacement of the M50 equivalent current dipole location in the left hemisphere. CONCLUSIONS: Cerebral lateralization and localization of the M50 distinguished patients with schizophrenia from normal subjects. These findings suggest the possibility of anatomical displacement and/or disturbed organization of the primary sensory cortex in schizophrenia.     

The dynamic changes in somatosensory input into the cortical motor area after damage to the somatosensory cortical zones in dogs

Dynamics of changes in the somatosensory evoked potentials (EPs) in the foreleg motor cortex (MI) was studied in awake dogs in chronic experiments following lesions of the forelimb area in the somatosensory cortex (SI) or in area 5. Cortical ablations reduced the amplitude of the primary positive-negative wave of the EPs in ipsilateral MI down to 28-63% of the original one. Within three weeks recovery of the EPs was insignificant or was absent at all. So there was no correlation between the EPs changes and spontaneous recovery of precise avoidance movements of the foreleg which takes place two weeks after local ablation of the SI. After area 5 lesion the EPs in the MI did not recover in spite of complete recovery of the EPs in the adjoining area of SI after its initial reduction down to 53%. The long-lasting EPs reduction was apparently a result of destruction of somatotopic sensory inputs from SI (or area 5) to MI, since EPs in MI of the intact hemisphere and in the hindlimb area of MI on the side of the lesion either did not change or recovered during one or two weeks.     

Face representation in the human secondary somatosensory cortex.

OBJECTIVE: To investigate the somatotopic organization of the facial skin area in the secondary somatosensory cortex (SII) in humans. METHODS: Somatosensory evoked magnetic fields following air-puff stimulation of 5 body sites, the foot, the lip and 3 points of the facial skin (forehead, cheek and mandibular angle point), were recorded. We focused on activities in SII following stimulation of these 5 sites and compared dipole locations among them. RESULTS: There was a clear somatotopic organization in SII with lip in the most lateral area, foot in the most medial area and face in an intermediate area close to the lip area. However, there was no significant difference of dipole localization in SII among the 3 areas of facial skin, similar to the overlapped somatotopic organization of facial skin areas in the primary somatosensory cortex in our previous study. CONCLUSIONS: The facial skin areas are considered to occupy a small area in SII with insufficient spatial separation to differentiate each area of facial skin even using magnetoencephalography which has a high spatial resolution. SIGNIFICANCE: This is the first systematic study of the activated regions in SII following stimulation of the facial skin.     

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.