Mikko Sams

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.

Seeing speech: visual information from lip movements modifies activity in the human auditory cortex

Neuromagnetic responses were recorded over the left hemisphere to find out in which cortical area the heard and seen speech are integrated. Auditory stimuli were Finnish/pa/syllables presented together with a videotaped face articulating either the concordant syllable/pa/(84% of stimuli, V = A) or the discordant syllable/ka/(16%, V not equal to A). In some subjects the probabilities were reversed. The subjects heard V not equal to A stimuli as/ta/ or ka. The magnetic responses to infrequent perceptions elicited a specific waveform which could be explained by activity in the supratemporal auditory cortex. The results show that visual information from articulatory movements has an entry into the auditory cortex.

Processing of changes in visual speech in the human auditory cortex

Seeing a talkers articulatory gestures may affect the observers auditory speech percept. Observing congruent articulatory gestures may enhance the recognition of speech sounds [J. Acoust. Soc. Am. 26 (1954) 212], whereas observing incongruent gestures may change the auditory percept phonetically as occurs in the McGurk effect [Nature 264 (1976) 746]. For example, simultaneous acoustic /ba/ and visual /ga/ are usually heard as /da/. We studied cortical processing of occasional changes in audiovisual and visual speech stimuli with magnetoencephalography. In the audiovisual experiment congruent (acoustic /iti/, visual /iti/) and incongruent (acoustic /ipi/, visual /iti/) audiovisual stimuli, which were both perceived as /iti/, were presented among congruent /ipi/ (acoustic /ipi/, visual /ipi/) stimuli. In the visual experiment only the visual components of these stimuli were presented. A visual change both in audiovisual and visual experiments activated supratemporal auditory cortices bilaterally. The auditory cortex activation to a visual change occurred later in the visual than in the audiovisual experiment, suggesting that interaction between modalities accelerates the detection of visual change in speech.

Face-Specific responses from the human inferior occipito-temporal cortex

Whole-head neuromagnetic responses were recorded from seven subjects to pictures of faces and to various control stimuli. Four subjects displayed signals specific to faces. The combination of functional information from magnetoencephalography and anatomical data from magnetic resonance images suggests that the face-specific activity was generated in the inferior occipitotemporal cortex. All four subjects showed the face-specific response in the right hemisphere, one of them also in the left. Our results, together with recent position emission tomography and lesion studies, suggest a right-hemisphere preponderance of face processing in the inferior occipitotemporal cortex.

Auditory evoked magnetic fields to tones and pseudowords in healthy children and adults.

Neuromagnetic responses to tones and pseudowords were measured with a 24-channel magnetometer in nine adults and in 23 children, the latter aged 0.3–15 years. Both stimulus types elicited substantially similar responses in all subjects. At 0.9-s interstimulus interval (ISI), the adult response was a stable P1 m-N1m-P2m-N2m sequence peaking at 50, 100, 200, and 250 ms, respectively. A biphasic Plm-N1,2m response with peaks at 100 and 260 ms occurred in children up to 12 years of age. At longer ISIs (1.2–2.4 s), an adult-type N1m response appeared in most children. N1 m amplitude suppression at short ISIs was stronger in children than in adults and may reflect a longer refractory period of the N1 generator neurons during early childhood than later in life. Peak latencies of PI m, N1 m, and N2m decreased with age, most rapidly <7 years of age. All deflections originated in nearby cortical areas within the posterior sylvian fissure, and may serve as functional landmarks for that anatomic area.

SEPARATE FINGER REPRESENTATIONS AT THE HUMAN SECOND SOMATOSENSORY CORTEX

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

Temporal integration in auditory sensory memory: neuromagnetic evidence

The cortical mechanisms of auditory sensory memory were investigated by analysis of neuromagnetic evoked responses. The major deflection of the auditory evoked field (N100m) appears to comprise an early posterior component (N100mP) and a late anterior component (N100mA) which is sensitive to temporal factors. When pairs of identical sounds are presented at intervals less that about 250 msec, the second sound evokes N100mA with enhanced amplitude at a latency of about 150 msec. We suggest that N100mA may index the activity of two distinct processes in auditory sensory memory. Its recovery cycle may reflect the activity of a memory trace which, according to previous studies, can retain processed information about an auditory sequence for about 10 sec. The enhancement effect may reflect the activity of a temporal integration process, whose time constant is such that sensation persists for 200-300 msec after stimulus offset, and so serves as a short memory store. Sound sequences falling within this window of integration seem to be coded holistically as unitary events.

Event-related potentials to repetition and change of auditory stimuli

The major intent of this study was to compare the role of stimulus repetition and change in the elicitation of the MMN, an ERP component specific to stimulus change, and N2b, usually partially overlapping the MMN when stimuli are attended. Event-related potentials were recorded in one set of conditions where subjects ignored the stimuli and read a book, and in another set of conditions where subjects counted stimuli designated as targets. Stimuli were delivered in 4 ways, the common feature between all these conditions being the occurrence of infrequent events at a probability of 0.20: (1) an oddball paradigm with 1 deviant, (2) an oddball paradigm with 2 deviants, each with a probability of 0.10, (3) a regular alternation of tones of 2 pitches where either of the 2 tones infrequently repeated (P = 0.20), and (4) a random presentation of tones of 5 different pitches, where any of the 5 tones infrequently repeated (P = 0.20). In the count conditions, the infrequent events were designated as targets. It was found that the MMN was elicited by stimulus change and not stimulus repetition in the ignore and count conditions, whereas the N2b was elicited by both stimulus changes and repetitions in the count conditions. It was also possible, in the count conditions, to disentangle the part of the late positive complex which is related to stimulus deviation and the part which is related to stimulus significance (target).

Seeing faces activates three separate areas outside the occipital visual cortex in man.

We have examined magnetic cortical responses of 15 healthy humans to 46 different pictures of faces. At least three areas outside the occipital visual cortex appeared to be involved in processing this input, 105-560 ms after the stimulus onset. The first active area was near the occipitotemporal junction, the second in the inferior parietal lobe, and the third in the middle temporal lobe. The source in the inferior parietal lobe was also activated by other simple and complex visual stimuli.

Visual attention modulates audiovisual speech perception

Speech perception is audiovisual, as demonstrated by the McGurk effect in which discrepant visual speech alters the auditory speech percept. We studied the role of visual attention in audiovisual speech perception by measuring the McGurk effect in two conditions. In the baseline condition, attention was focused on the talking face. In the distracted attention condition, subjects ignored the face and attended to a visual distractor, which was a leaf moving across the face. The McGurk effect was weaker in the latter condition, indicating that visual attention modulated audiovisual speech perception. This modulation may occur at an early, unisensory processing stage, or it may be due to changes at the stage where auditory and visual information is integrated. We investigated this issue by conventional statistical testing, and by fitting the Fuzzy Logical Model of Perception (Massaro, 1998) to the results. The two methods suggested different interpretations, revealing a paradox in the current methods of analysis.