Processing of novel sounds and frequency changes in the human auditory cortex: Magnetoencephalographic recordings

Whole-head magnetoencephalographic (MEG) responses to repeating standard tones and to infrequent slightly higher deviant tones and complex novel sounds were recorded together with event-related brain potentials (ERPs). Deviant tones and novel sounds elicited the mismatch negativity (MMN) component of the ERP and its MEG counterpart (MMNm) both when the auditory stimuli were attended to and when they were ignored. MMNm generators were located bilateral to the superior planes of the temporal lobes where preattentive auditory discrimination appears to occur. A subsequent positive P3a component was elicited by deviant tones and with a larger amplitude by novel sounds even when the sounds were to be ignored. Source localization for the MEG counterpart of P3a (P3am) suggested that the auditory cortex in the superior temporal plane is involved in the neural network of involuntary attention switching to changes in the acoustic environment.     

Effects of voluntary hyperventilation on cortical sensory responses Electroencephalographic and magnetoencephalographic studies

 It is well established that voluntary hyperventilation (HV) slows down electroencephalographic (EEG) rhythms. Little information is available, however, on the effects of HV on cortical responses elicited by sensory stimulation. In the present study, we recorded auditory evoked potentials (AEPs) and magnetic fields (AEFs), and somatosensory evoked magnetic fields (SEFs) from healthy subjects before, during, and after a 3- to 5-min period of voluntary HV. The effectiveness of HV was verified by measuring the end-tidal CO₂ levels. Long-latency (100–200 ms) AEPs and long-latency AEFs originating at the supratemporal auditory cortex, as well as long-latency SEFs from the primary somatosensory cortex (SI) and from the opercular somatosensory cortex (OC), were all reduced during HV. The short-latency SEFs from SI were clearly less modified, there being, however, a slight reduction of the earliest cortical excitatory response, the N20m deflection. A middle-latency SEF deflection from SI at about 60 ms (P60 m) was slightly increased. For AEFs and SEFs, the center-of-gravity locations of the activated neuronal populations were not changed during HV. All amplitude changes returned to baseline levels within 10 min after the end of HV. The AEPs were not altered when the subjects breathed 5% CO₂ in air in a hyperventilation-like manner, which prevented the development of hypocapnia. We conclude that moderate HV suppresses long-latency evoked responses from the primary projection cortices, while the early responses are less reduced. The reduction of long-latency responses is probably mediated by hypocapnia rather than by other nonspecific effects of HV. It is suggested that increased neuronal excitability caused by HV-induced hypocapnia leads to spontaneous and/or asynchronous firing of cortical neurones, which in turn reduces stimulus-locked synaptic events. Received: 14 October 1997 / Accepted: 28 October 1998     

Context modulates processing of speech sounds in the right auditory cortex of human subjects

Functionally, the cerebellum is not only involved in motor control but is also thought to influence higher cognitive function including language. Anatomical data would suggest crossed reciprocal connections between the cerebellum and higher order cortical association areas. In the following study, one left- and one right-handed female volunteer underwent functional magnetic resonance imaging in a conventional block design. Regions of activation were detected after performance of a silent verbal fluency task inside the scanner. In the right-handed volunteer we found an activation of the left fronto-parietal cortex and the right cerebellar hemisphere, while in the left-handed volunteer the activation was seen in the right fronto-parieto-temporal cortex and the left cerebellar hemisphere. These initial results demonstrate that cerebellar activation is contralateral to the activation of the frontal cortex even under conditions of different language dominance. They provide evidence for the hypothesis of a lateralized organization of the cerebellum crossed to the cerebral hemispheres in supporting higher cognitive function.     

Methodology for Combined TMS and EEG

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.     

Spatiotemporal activity of a cortical network for processing visual motion revealed by MEG and fMRI

A sudden change in the direction of motion is a particularly salient and relevant feature of visual information. Extensive research has identified cortical areas responsive to visual motion and characterized their sensitivity to different features of motion, such as directional specificity. However, relatively little is known about responses to sudden changes in direction. Electrophysiological data from animals and functional imaging data from humans suggest a number of brain areas responsive to motion, presumably working as a network. Temporal patterns of activity allow the same network to process information in different ways. The present study in humans sought to determine which motion-sensitive areas are involved in processing changes in the direction of motion and to characterize the temporal patterns of processing within this network of brain regions. To accomplish this, we used both magnetoencephalography (MEG) and functional magnetic resonance imaging (fMRI). The fMRI data were used as supplementary information in the localization of MEG sources. The change in the direction of visual motion was found to activate a number of areas, each displaying a different temporal behavior. The fMRI revealed motion-related activity in areas MT+ (the human homologue of monkey middle temporal area and possibly also other motion sensitive areas next to MT), a region near the posterior end of the superior temporal sulcus (pSTS), V3A, and V1/V2. The MEG data suggested additional frontal sources. An equivalent dipole model for the generators of MEG signals indicated activity in MT+, starting at 130 ms and peaking at 170 ms after the reversal of the direction of motion, and then again at approximately 260 ms. Frontal activity began 0-20 ms later than in MT+, and peaked approximately 180 ms. Both pSTS and FEF+ showed long-duration activity continuing over the latency range of 200-400 ms. MEG responses in the region of V3A and V1/V2 were relatively small, and peaked at longer latencies than the initial peak in MT+. These data revealed characteristic patterns of activity in this cortical network for processing sudden changes in the direction of visual motion.     

Projecting out muscle artifacts from TMS-evoked EEG

Transcranial magnetic stimulation combined with electroencephalography is a powerful tool for probing cortical excitability and connectivity; we can perturb one brain area and study the reactions at the stimulated and interconnected sites. When stimulating areas near cranial muscles, their activation produces a large artifact in the electroencephalographic signal, lasting tens of milliseconds and masking the early brain signals. We present an artifact removal method based on projecting out the topographic patterns of the muscle activity. Although the brain and muscle components overlap both temporally and spectrally, the fact that muscle activity is present also at frequencies higher than 100 Hz, while brain signal is mostly restricted to frequencies lower than that, allows us to study the high-frequency muscle activity without brain contribution. We determined the muscle activity topographies from data highpass-filtered at a 100-Hz cutoff frequency using principal component analysis. Projecting out the topographies of the principal components which explain most of the variance of the high-frequency data reduces not only the high-frequency activity but also the low-frequency muscle contribution, because the topography produced by a muscle source can be expected to be the same regardless of the frequency. The method greatly reduced the muscle artifact evoked by stimulation of Broca's area, while a significant brain signal contribution remained. Improvement in the signal-to-artifact ratio, defined as the relative amplitudes of brain signals peaking after 50 ms and the first artifact deflection, was of the order of 10-100 depending on the number of projections. The presented artifact removal method enables one to study the cortical state when stimulating areas near the cranial muscles.     

A novel mechanism for evoked responses in the human brain

Magnetoencephalographic and electroencephalographic evoked responses are primary real-time objective measures of cognitive and perceptual processes in the human brain. Two mechanisms (additive activity and phase reset) have been debated and considered as the only possible explanations for evoked responses. Here we present theoretical and empirical evidence of a third mechanism contributing to the generation of evoked responses. Interestingly, this mechanism can be deduced entirely from the characteristics of spontaneous oscillations in the absence of stimuli. We show that the amplitude fluctuations of neuronal alpha oscillations at rest are associated with changes in the mean value of ongoing activity in magnetoencephalography, a phenomenon that we term baseline shifts associated with alpha oscillations. When stimuli modulate the amplitude of alpha oscillations, baseline shifts become the basis of a novel mechanism for the generation of evoked responses; the averaging of several trials leads to a cancellation of the oscillatory component but the baseline shift remains, which gives rise to an evoked response. We propose that the presence of baseline shifts associated with alpha oscillations can be explained by the asymmetric flow of inward and outward neuronal currents related to the generation of alpha oscillations. Our findings are relevant to the vast majority of electroencephalographic and magnetoencephalographic studies involving perceptual, cognitive and motor activity.     

The interplay of lorazepam-induced brain oscillations: microstructural electromagnetic study.

OBJECTIVE: The effects on cortical rhythms of a single-dose (30 microg/kg) administration of the GABAA agonist lorazepam were examined in a randomized, double-blind, cross-over, placebo-controlled study with 8 healthy volunteers using simultaneous electroencephalography (EEG) and magnetoencephalography (MEG). METHODS: The oscillations were assessed by means of adaptive classification of short-term spectral patterns. RESULTS: Lorazepam (a) decreased the percentage of EEG/MEG segments with fast-theta, delta-alpha, fast-theta-alpha and alpha activity and increased percentage of EEG/MEG segments with delta, delta-slow-theta, delta-beta, slow-theta and polyrhythmic activity; (b) decreased diversity of EEG/MEG signals (in terms of spectral patterns) and increased the general instability of the signal; (c) increased stabilization periods of the spectral patterns (reduced brain information processing); (d) maintained larger maximum periods of temporal stabilization for delta, slow-theta, delta-slow-theta, delta-beta and polyrhythmic activity (in terms of spectral patterns); (e) did not increase power in the independent beta rhythm. CONCLUSIONS: Lorazepam caused significant reorganization of the EEG/MEG microstructure. These results suggest also that adaptive classification analysis of single short-term spectral patterns may provide additional information to conventional spectral analyses.     

Ipsi- and contralateral EEG reactions to transcranial magnetic stimulation.

OBJECTIVES: Transcranial magnetic stimulation (TMS) and high-resolution electroencephalography (EEG) were used to study the spreading of cortical activation in 6 healthy volunteers. METHODS: Five locations in the left sensorimotor cortex (within 3cm(2)) were stimulated magnetically, while EEG was recorded with 60 scalp electrodes. A frameless stereotactic method was applied to determine the anatomic locus of stimulation and to superimpose the results on magnetic resonance images. Scalp potential and cortical current-density distributions were derived from averaged electroencephalographic (EEG) data. RESULTS: The maxima of the ipsilateral activation were detected at the gyrus precentralis, gyrus supramarginalis, and lobulus parietalis superior, depending on the subject. Activation over the contralateral cortex was observed in all subjects, appearing at 22plus minus2ms (range 17--28); the maxima were located at the gyrus precentralis, gyrus frontalis superior, and the lobulus parietalis inferior. Contralateral EEG waveforms showed consistent changes when different sites were stimulated: stimulation of the two most medial points evoked the smallest responses fronto-parietally. CONCLUSIONS: With the combination of TMS, EEG, and magnetic resonance imaging, an adequate spatiotemporal resolution may be achieved for tracing the intra- and interhemispheric spread of activation in the cortex caused by a magnetic pulse.     

Methods for analysis of brain connectivity: An IFCN-sponsored review

The goal of this paper is to examine existing methods to study the “Human Brain Connectome” with a specific focus on the neurophysiological ones. In recent years, a new approach has been developed to evaluate the anatomical and functional organization of the human brain: the aim of this promising multimodality effort is to identify and classify neuronal networks with a number of neurobiologically meaningful and easily computable measures to create its connectome. By defining anatomical and functional connections of brain regions on the same map through an integrated approach, comprising both modern neurophysiological and neuroimaging (i.e. flow/metabolic) brain-mapping techniques, network analysis becomes a powerful tool for exploring structural–functional connectivity mechanisms and for revealing etiological relationships that link connectivity abnormalities to neuropsychiatric disorders. Following a recent IFCN-endorsed meeting, a panel of international experts was selected to produce this current state-of-art document, which covers the available knowledge on anatomical and functional connectivity, including the most commonly used structural and functional MRI, EEG, MEG and non-invasive brain stimulation techniques and measures of local and global brain connectivity.