Magnetoencephalography(MEG) is a noninvasive technique for investigating neuronal activity in the living human brairi. The time resolution of the method is better than 1 ms and the spatial discrimination is, under favorable circumstances, 2 -3 mm for sources in the cerebral cortex. In MEG studies, the weak 10 fT -1 pT magnetic fields produced by electric currents fiowing in neurons are measured with multichannel SQUID {superconducting quantum interference device) gradiometers. The sites in the cerebral cortex that are activated by a stimulus can be found from the detected magnetic-field distribution, provided that appropriate assumptions about the source render the solution of the inverse problem unique. Many interesting properties of the working human brain can be studied, including spontaneous activity and signal processing following external stimuli. For clinical purposes, determination of the locations of epileptic foci is of interest. The authors begiri with a general introduction and a short discussion of the neural basis of MEG. The mathematical theory of the method is then explained in detail, followed by a thorough description of MEG instrumentation, data analysis, and practical construction of multi-SQUID devices. Finally, several MEG experiments performed in the authors laboratory are described, covering studies of evoked responses and of spontaneous activity in both healthy and diseased brains. Many MEG studies by other groups are discussed briefiy as well.

The authors have applied estimation theory to the problem of determining primary current distributions from measured neuromagnetic fields. In this procedure, essentially nothing is assumed about the source currents, except that they are spatially restricted to a certain region. Simulation experiments show that the results can describe the structure of the current flow fairly well. By increasing the number of measurements, the estimate can be made more localised. The current distributions may be also used as an interpolation and an extrapolation for the measured field patterns.

There is considerable debate about whether the early processing of sounds depends on whether they form part of speech. Proponents of such speech specificity postulate the existence of language-dependent memory traces, which are activated in the processing of speech but not when equally complex, acoustic non-speech stimuli are processed. Here we report the existence of these traces in the human brain. We presented to Finnish subjects the Finnish phoneme prototype /e/ as the frequent stimulus, and other Finnish phoneme prototypes or a non-prototype (the Estonian prototype /õ/) as the infrequent stimulus. We found that the brain's automatic change-detection response, reflected electrically as the mismatch negativity (MMN), was enhanced when the infrequent, deviant stimulus was a prototype (the Finnish /ö/) relative to when it was a non-prototype (the Estonian /õ/). These phonemic traces, revealed by MMN, are language-specific, as /õ/ caused enhancement of MMN in Estonians. Whole-head magnetic recordings located the source of this native-language, phoneme-related response enhancement, and thus the language-specific memory traces, in the auditory cortex of the left hemisphere.

Motor and visual cortices of normal volunteers were activated by transcranial magnetic stimulation. The electrical brain activity resulting from the brief electromagnetic pulse was recorded with high-resolution electroencephalography (HR-EEG) and located using inversion algorithms. The stimulation of the left sensorimotor hand area elicited an immediate response at the stimulated site. The activation had spread to adjacent ipsilateral motor areas within 5-10 ms and to homologous regions in the opposite hemisphere within 20 ms. Similar activation patterns were generated by magnetic stimulation of the visual cortex. This new non-invasive method provides direct information about cortical reactivity and area-to-area neuronal connections.

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Transcranial magnetic stimulation (TMS) was applied to motor areas in the left language-dominant hemisphere while right-handed human subjects made lexical decisions on words related to actions. Response times to words referring to leg actions (e.g. kick) were compared with those to words referring to movements involving the arms and hands (e.g. pick). TMS of hand and leg areas influenced the processing of arm and leg words differentially, as documented by a significant interaction of the factors Stimulation site and Word category. Arm area TMS led to faster arm than leg word responses and the reverse effect, faster lexical decisions on leg than arm words, was present when TMS was applied to leg areas. TMS-related differences between word categories were not seen in control conditions, when TMS was applied to hand and leg areas in the right hemisphere and during sham stimulation. Our results show that the left hemispheric cortical systems for language and action are linked to each other in a category-specific manner and that activation in motor and premotor areas can influence the processing of specific kinds of words semantically related to arm or leg actions. By demonstrating specific functional links between action and language systems during lexical processing, these results call into question modular theories of language and motor functions and provide evidence that the two systems interact in the processing of meaningful information about language and action.

Life or death in hostile environments depends crucially on one's ability to detect and gate novel sounds to awareness, such as that of a twig cracking under the paw of a stalking predator in a noisy jungle. Two distinct auditory cortex processes have been thought to underlie this phenomenon: (i) attenuation of the so-called N1 response with repeated stimulation and (ii) elicitation of a mismatch negativity response (MMN) by changes in repetitive aspects of auditory stimulation. This division has been based on previous studies suggesting that, unlike for the N1, repetitive ''standard'' stimuli preceding a physically different ''novel'' stimulus constitute a prerequisite to MMN elicitation, and that the source loci of MMN and N1 are different. Contradicting these findings, our combined electromagnetic, hemodynamic, and psychophysical data indicate that the MMN is generated as a result of differential adaptation of anterior and posterior auditory cortex N1 sources by preceding auditory stimulation. Early (Ϸ85 ms) neural activity within posterior auditory cortex is adapted as sound novelty decreases. This alters the center of gravity of electromagnetic N1 source activity, creating an illusory difference between N1 and MMN source loci when estimated by using equivalent current dipole fits. Further, our electroencephalography data show a robust MMN after a single standard event when the interval between two consecutive novel sounds is kept invariant. Our converging findings suggest that transient adaptation of feature-specific neurons within human posterior auditory cortex filters superfluous sounds from entering one's awareness.

The combination of transcranial magnetic stimulation (TMS) with simultaneous electroencephalography (EEG) provides us the possibility to non-invasively probe the brain's excitability, time-resolved connectivity and instantaneous state. Early attempts to combine TMS and EEG suffered from the huge electromagnetic artifacts seen in EEG as a result of the electric field induced by the stimulus pulses. To deal with this problem, TMS-compatible EEG systems have been developed. However, even with amplifiers that are either immune to or recover quickly from the pulse, great challenges remain. Artifacts may arise from the movement of electrodes, from muscles activated by the pulse, from eye movements, from electrode polarization, or from brain responses evoked by the coil click. With careful precautions, many of these problems can be avoided. The remaining artifacts can be usually reduced by filtering, but control experiments are often needed to make sure that the measured signals actually originate in the brain. Several studies have shown the power of TMS-EEG by giving us valuable information about the excitability or connectivity of the brain.