Late pain-related magnetic fields and electric potentials evoked by intracutaneous electric finger stimulation

Surface magnetic and electric recordings were used to localize the sources of late pain-related magnetic fields and electric potentials, evoked by painful intracutaneous electric finger stimulation. We find that the source of the P90m component of the evoked magnetic field lies in the finger area of the primary somatosensory cortex; the sources of the N150m and P250m are found to reside in the frontal operculum. These findings are unexpected from the evoked electric potential data, which suggest a central location for these sources. We also note that the interpretation of the electric data was confounded by the presence of an alpha-like oscillation, which overlapped many components of the evoked potential.     

Independent Components of Neural Activity Carry Information on Individual Populations

Local field potential (LFP), the low-frequency part of the potential recorded extracellularly in the brain, reflects neural activity at the population level. The interpretation of LFP is complicated because it can mix activity from remote cells, on the order of millimeters from the electrode. To understand better the relation between the recordings and the local activity of cells we used a large-scale network thalamocortical model to compute simultaneous LFP, transmembrane currents, and spiking activity. We used this model to study the information contained in independent components obtained from the reconstructed Current Source Density (CSD), which smooths transmembrane currents, decomposed further with Independent Component Analysis (ICA). We found that the three most robust components matched well the activity of two dominating cell populations: superior pyramidal cells in layer 2/3 (rhythmic spiking) and tufted pyramids from layer 5 (intrinsically bursting). The pyramidal population from layer 2/3 could not be well described as a product of spatial profile and temporal activation, but by a sum of two such products which we recovered in two of the ICA components in our analysis, which correspond to the two first principal components of PCA decomposition of layer 2/3 population activity. At low noise one more cell population could be discerned but it is unlikely that it could be recovered in experiment given typical noise ranges.     

Cross-population coupling of neural activity based on Gaussian process current source densities

Because local field potentials (LFPs) arise from multiple sources in different spatial locations, they do not easily reveal coordinated activity across neural populations on a trial-to-trial basis. As we show here, however, once disparate source signals are decoupled, their trial-to-trial fluctuations become more accessible, and cross-population correlations become more apparent. To decouple sources we introduce a general framework for estimation of current source densities (CSDs). In this framework, the set of LFPs result from noise being added to the transform of the CSD by a biophysical forward model, while the CSD is considered to be the sum of a zero-mean, stationary, spatiotemporal Gaussian process, having fast and slow components, and a mean function, which is the sum of multiple time-varying functions distributed across space, each varying across trials. We derived biophysical forward models relevant to the data we analyzed. In simulation studies this approach improved identification of source signals compared to existing CSD estimation methods. Using data recorded from primate auditory cortex, we analyzed trial-to-trial fluctuations in both steady-state and task-evoked signals. We found cortical layer-specific phase coupling between two probes and showed that the same analysis applied directly to LFPs did not recover these patterns. We also found task-evoked CSDs to be correlated across probes, at specific cortical depths. Using data from Neuropixels probes in mouse visual areas, we again found evidence for depth-specific phase coupling of primary visual cortex and lateromedial area based on the CSDs.     

Answering six questions in extracting children��s mismatch negativity through combining wavelet decomposition and independent component analysis

This study combines wavelet decomposition and independent component analysis (ICA) to extract mismatch negativity (MMN) from electroencephalography (EEG) recordings. As MMN is a small event-related potential (ERP), a systematic ICA based approach is designed, exploiting MMN’s temporal, frequency and spatial information. Moreover, this study answers which type of EEG recordings is more appropriate for ICA to extract MMN, what kind of the preprocessing is beneficial for ICA decomposition, which algorithm of ICA can be chosen to decompose EEG recordings under the selected type, how to determine the desired independent component extracted by ICA, how to improve the accuracy of the back projection of the selected independent component in the electrode field, and what can be finally obtained with the application of ICA. Results showed that the proposed method extracted MMN with better properties than those estimated by difference wave only using temporal information or ICA only using spatial information. The better properties mean that the deviant with larger magnitude of deviance to repeated stimuli in the oddball paradigm can elicit MMN with larger peak amplitude and shorter latency. As other ERPs also have the similar information exploited here, the proposed method can be used to study other ERPs.     

Empirical mode decomposition of field potentials from macaque V4 in visual spatial attention

Empirical mode decomposition (EMD) has recently been introduced as a local and fully data-driven technique for the analysis of non-stationary time-series. It allows the frequency and amplitude of a time-series to be evaluated with excellent time resolution. In this article we consider the application of EMD to the analysis of neuronal activity in visual cortical area V4 of a macaque monkey performing a visual spatial attention task. We show that, by virtue of EMD, field potentials can be resolved into a sum of intrinsic components with different degrees of oscillatory content. Low-frequency components in single-trial recordings contribute to the average visual evoked potential (AVEP), whereas high-frequency components do not, but are identified as gamma-band (30–90 Hz) oscillations. The magnitude of time-varying gamma activity is shown to be enhanced when the monkey attends to a visual stimulus as compared to when it is not attending to the same stimulus. Comparison with Fourier analysis shows that EMD may offer better temporal and frequency resolution. These results support the idea that the magnitude of gamma activity reflects the modulation of V4 neurons by visual spatial attention. EMD, coupled with instantaneous frequency analysis, is demonstrated to be a useful technique for the analysis of neurobiological time-series.     

Reconstructing Coherent Networks from Electroencephalography and Magnetoencephalography with Reduced Contamination from Volume Conduction or Magnetic Field Spread

Volume conduction (VC) and magnetic field spread (MFS) induce spurious correlations between EEG/MEG sensors, such that the estimation of functional networks from scalp recordings is inaccurate. Imaginary coherency [1] reduces VC/MFS artefacts between sensors by assuming that instantaneous interactions are caused predominantly by VC/MFS and do not contribute to the imaginary part of the cross-spectral densities (CSDs). We propose an adaptation of the dynamic imaging of coherent sources (DICS) [2] - a method for reconstructing the CSDs between sources, and subsequently inferring functional connectivity based on coherences between those sources. Firstly, we reformulate the principle of imaginary coherency by performing an eigenvector decomposition of the imaginary part of the CSD to estimate the power that only contributes to the non-zero phase-lagged (NZPL) interactions. Secondly, we construct an NZPL-optimised spatial filter with two a priori assumptions: (1) that only NZPL interactions exist at the source level and (2) the NZPL CSD at the sensor level is a good approximation of the projected source NZPL CSDs. We compare the performance of the NZPL method to the standard method by reconstructing a coherent network from simulated EEG/MEG recordings. We demonstrate that, as long as there are phase differences between the sources, the NZPL method reliably detects the underlying networks from EEG and MEG. We show that the method is also robust to very small phase lags, noise from phase jitter, and is less sensitive to regularisation parameters. The method is applied to a human dataset to infer parts of a coherent network underpinning face recognition.     

Dynamics of the EEG spectral density in the θ, α, and β bands in the visual <Emphasis Type="Italic">Go</Emphasis>/<Emphasis Type="Italic">NoGo</Emphasis> task

The study was aimed at analyzing event-related desynchronization/synchronization (ERD/ERS) in 19-channel EEGs recorded in 329 healthy subjects in the course of a Go/NoGo task. Three methods were tested: reference, current source density (CSD), and group decomposition by independent component analysis (ICA). A comparison of the three data sets showed that the ICA method better reflects the local features of the brain responses in the θ, α, and β ranges. The functional significance of the group ICA components is discussed.     

BICAR: A New Algorithm for Multiresolution Spatiotemporal Data Fusion

We introduce a method for spatiotemporal data fusion and demonstrate its performance on three constructed data sets: one entirely simulated, one with temporal speech signals and simulated spatial images, and another with recorded music time series and astronomical images defining the spatial patterns. Each case study is constructed to present specific challenges to test the method and demonstrate its capabilities. Our algorithm, BICAR (Bidirectional Independent Component Averaged Representation), is based on independent component analysis (ICA) and extracts pairs of temporal and spatial sources from two data matrices with arbitrarily different spatiotemporal resolution. We pair the temporal and spatial sources using a physical transfer function that connects the dynamics of the two. BICAR produces a hierarchy of sources ranked according to reproducibility; we show that sources which are more reproducible are more similar to true (known) sources. BICAR is robust to added noise, even in a “worst case” scenario where all physical sources are equally noisy. BICAR is also relatively robust to misspecification of the transfer function. BICAR holds promise as a useful data-driven assimilation method in neuroscience, earth science, astronomy, and other signal processing domains.     

GABAA Receptor in the Thalamic Specific Relay System Contributes to the Propofol-Induced Somatosensory Cortical Suppression in Rat

Interaction with the gamma-aminobutyric-acid-type-A (GABA_A) receptors is recognized as an important component of the mechanism of propofol, a sedative-hypnotic drug commonly used as anesthetic. However the contribution of GABA_A receptors to the central nervous system suppression is still not well understood, especially in the thalamocortical network. In the present study, we investigated if intracerebral injection of bicuculline (a GABA_A receptor antagonist) into the thalamus ventral posteromedial nucleus (VPM, a thalamus specific relay nuclei that innervated S1 mostly) could reverse propofol-induced cortical suppression, through recording the changes of both spontaneous and somatosensory neural activities in rat’s somatosensory cortex (S1). We found that after injection of bicuculline into VPM, significant increase of neural activities were observed in all bands of local field potentials (total band, 182±6%), while the amplitude of all components in somatosensory evoked potentials were also increased (negative, 121±9% and positive, 124±6%).These data support that the potentiation of GABA_A receptor-mediated synaptic inhibition in a thalamic specific relay system seems to play a crucial role in propofol-induced cortical suppression in the somatosensory cortex of rats.     

Brain-stem somatosensory dysfunction in a case of long-standing left hemispherectomy with removal of the left thalamus: a nasopharyngeal and scalp SEP study

We have studied median nerve somatosensory evoked potentials (SEPs) in a patient who had undergone early surgical removal of the left cerebral hemisphere and left thalamus. Stimulation of the right side evoked normal latency P9, P11 and P13 potentials at scalp as well as at nasopharyngeal (NP) leads, while P14 and N18 potentials were absent. These SEP abnormalities, that have been described previously in cervico-medullary lesions and in comatose patients with upper brain-stem involvement, suggest that in our patient the removal of the left thalamus has caused retrograde degeneration of the cuneate-thalamic projections. Moreover, this study confirms that P13 and P14 potentials have different generators.     

Cross-trial correlation analysis of evoked potentials reveals arousal-related attenuation of thalamo-cortical coupling

We describe a computational method for assessing functional connectivity in sensory neuronal networks. The method, which we term cross-trial correlation, can be applied to signals representing local field potentials (LFPs) evoked by sensory stimulations and utilizes their trial-to-trial variability. A set of single trial samples of a given post-stimulus latency from consecutive evoked potentials (EPs) recorded at a given site is correlated with such sets for all other latencies and recording sites. The results of this computation reveal how neuronal activities at various sites and latencies correspond to activation of other sites at other latencies. The method was used to investigate the functional connectivity of thalamo-cortical network of somatosensory system in behaving rats at two levels of alertness: habituated and aroused. We analyzed potentials evoked by vibrissal deflections recorded simultaneously from the ventrobasal thalamus and barrel cortex. The cross-trial correlation analysis applied to the early post-stimulus period (<25 ms) showed that the magnitude of the population spike recorded in the thalamus at 5 ms post-stimulus correlated with the cortical activation at 6–13 ms post-stimulus. This correlation value was reduced at 6–9 ms, i.e. at early postsynaptic cortical response, with increased level of the animals’ arousal. Similarly, the aroused state diminished positive thalamo-cortical correlation for subsequent early EP waves, whereas the efficacy of an indirect cortico-fugal inhibition (over 15 ms) did not change significantly. Thus we were able to characterize the state related changes of functional connections within the thalamo-cortical network of behaving animals.     

Somatotopy of human hand somatosensory cortex as studied in scalp EEG

We recorded somatosensory evoked potentials (SEPs) in scalp EEGs during stimulation of the median nerve, the ulnar nerve and the individual digits in 3 normal subjects and in 1 epilepsy patients. In this patient we also measured SEPs from chronically indwelling subdural grid electrodes during electrocorticography (ECoG). We applied dipole modelling techniques to study the 3-dimensional intracerebral locations and time activities of the neuronal sources underlying stimulation of different peripheral receptive fields. The sources underlying median nerve SEPs were located an average of 10.8 mm lateral inferior to those underlying ulnar nerve SEPs. Digit SEP sources showed a somatotopic arrangement from lateral inferior to medial superior in the order thumb, index finger, middle finger, ring finger and little finger, with some overlap or reversal for adjacent digits. The average distance between thumb and little finger was 12.5 mm. Thumb, index finger and middle finger were clustered around median nerve cortical representation, whereas ring finger and little finger were arranged around ulnar nerve cortex. In the epilepsy patient, the source localizations obtained in scalp EEGs showed good agreement with those on ECoGs. We conclude that SEPs recorded in scalp EEGs can be used to study functional topography of human somatosensory cortex non-invasively.     

Current source-density analysis in the mushroom bodies of the honeybee (Apis mellifera carnica)

Summary Information processing in the mushroom bodies which are an important part of most invertebrate central nervous systems was analysed by extracellular electrophysiological techniques. The mushroom bodies consist of layers of parallel intrinsic neurons which make synaptic contact with extrinsic input and output neurons. The intrinsic neurons (approximately 170,000/mushroom body) have very small axon diameters (0.1–1 m) which makes it difficult to record their activity intracellularly. In order to analyse the functional properties of this neuropil field potentials were measured extracellularly.Series of averaged evoked potentials (AEPs) were recorded along electrode tracks at consecutive depth intervals in different parts of the mushroom bodies of the bee. These potentials were elicited by olfactory, mechanical and visual stimuli.In order to locate the synaptic areas generating these potentials, current source-densities (CSD) were calculated using the consecutively measured evoked potentials. The conductivities of the extracellular space along the electrode tracks in the pedunculus and calyx and in part of the alpha-lobe of the mushroom bodies were found to be constant.The CSD analysis reveals a complex pattern of source-sink distributions in the mushroom bodies. There is a high degree of correlation between current sinks and sources detected by CSD analysis and the morphological distribution of neurons.The CSD analysis shows that the inputs and outputs of the mushroom bodies involve multimodal synaptic interactions, whereas information processing in the intrinsic Kenyon-cells is limited to sensory inputs from the antenna.Comparison of the electrophysiological with the histological results shows that the intrinsic cells of the mushroom bodies are physiologically not a homogeneous group as is often proposed. Among the intrinsic neurons clearly defined areas of current sources and sinks can be identified and attributed to Kenyon-cells in different layers.Abbreviations AEP averaged evoked potentials - AGT antennoglomerular tract - CSD current source-density - PCT antennoglomerular tract     

Changes in somatosensory evoked potentials following an experimental focal ischaemic lesion in thalamus

Experiments have been performed to produce localized thalamic ischaemia in baboons anaesthetised with alpha-chloralose. Somatosensory evoked potentials to median nerve stimulation were recorded in the medial lemniscus. VPL of thalamus and the primary somatosensory cortex. Local blood flow was also recorded by the hydrogen clearance technique in these regions. The early potential recorded in thalamus has been shown to be generated from 3 sources: (i) a positivity generated outside the VPL, (ii) local wavelets, most likely from synaptic activity close to the recording electrode, and (iii) a local overall negativity. The first of these potentials alone remains after thalamic ischaemia. It arises below the level of the thalamus, being very likely generated by the afferent volley in the medial lemniscus, and is seen in the surface-recorded response as the early component P8 (corresponding to P15 in the human).     

Multichannel visual evoked potentials in migraine

Multichannel recordings of visual evoked potentials (VEPs) have proved to be useful in the evaluation of visual field defects. We studied the topographic distribution of transient VEPs in 15 migraine patients (8 with visual aura and 7 without) and 15 age-matched controls during the migraine-free interval. All the subjects included in the study had normal visual fields. VEPs were recorded from 9 electrodes placed on the posterior scalp. Stimuli were full-field and hemifield reversing square wave grating patterns of medium spatial frequency (4 c/deg). The groups did not show significant differences in latencies and amplitudes of the major components (N70, P100) recorded from the midline. However, migraine patients with visual hemianopic aura showed definite asymmetries in the VEP amplitude distribution. Significantly reduced, absent or polarity-invered VEP responses were recorded ipsilateral to the side of the prodromic visual symptoms. Direct comparison of affected and unaffected hemispheres by partial field stimulation confirmed these findings. According to the VEP cortical generator theory, these abnormalities suggest a functional anomaly consistent with the clinical syndrome and detectable also in the migraine-free interval. None of the migraine patients without aura or the controls showed VEP amplitude asymmetries. We conclude that multichannel VEP recordings may discriminate between different subtypes of migraine and contribute important physiopathological information to the study of this disease.     

Influence of acute cerebellar lesions on somatosensory evoked potentials (SEPs) in cats.

We studied the effect of acute unilateral cerebellar lesions on the cerebello-thalamo-cortical projection in cats. The lesions were classified into two groups according to their extent. In group I the lesion only covered the cerebellar cortex, while in group II both the cerebellar cortex and deep cerebellar nuclei were removed. Early (short-latency) and late (long-latency) waves, evoked by an electrical stimulation of a forelimb, were collected contralateral to the stimulated leg hemisphere. Pre- and postsurgery recordings from primary and non-primary (motor and parietal) cortices were compared. Cerebellar impairment had a strong influence on discharges of all the considered cortical areas. Early non-primary and primary responses increased in group I and remained unchanged in group II. Late somatosensory evoked potentials components were suppressed in both groups. An inhibitory influence of the cerebellar cortex on the thalamo-cortical projection was confirmed. Changes within the primary sensory cortex may suggest an engagement of that area in the compensation process of cerebellar dysfunction shortly after cerebellar lesion. An alteration in the unaffected hemisphere activation indicate that the spino-cerebellar and cerebello-cortical inputs, responsible for somatosensory evoked potentials generation, are regulated through contralateral and ipsilateral pathways. These pathways are unmasked by cerebellar lesion.     

The second somatosensory region of the cat thalamus

Recording focal evoked potentials showed that the second somatosensory region of the cat thalamus is located in the posterolateral part of the ventral posteromedial nucleus. The zone of representation of the hindlimb lies somewhat laterally, posteriorly, and superiorly to the zone of the forelimb. The zone of representation of the mouth lies more medially still. Latent periods of responses arising in both regions of the thalamus to cutaneous stimulation of the contralateral limbs are about equal. Neurons of the second somatosensory region of the thalamus send projections to cortical area S2. It can be concluded from the results that formation of the structure of double representation of the somatosensory systems in the cat is completed at the thalamic level.     

Synaptic mechanisms underlying sparse coding of active touch

Sensory information is actively gathered by animals, but the synaptic mechanisms driving neuronal circuit function during active sensory processing are poorly understood. Here, we investigated the synaptically driven membrane potential dynamics during active whisker sensation using whole-cell recordings from layer 2/3 pyramidal neurons in the primary somatosensory barrel cortex of behaving mice. Although whisker contact with an object evoked rapid depolarization in all neurons, these touch responses only drove action potentials in ～10% of the cells. Such sparse coding was ensured by cell-specific reversal potentials of the touch-evoked response that were hyperpolarized relative to action potential threshold for most neurons. Intercontact interval profoundly influenced touch-evoked postsynaptic potentials, interestingly without affecting the peak membrane potential of the touch response. Dual whole-cell recordings indicated highly correlated membrane potential dynamics during active touch. Sparse action potential firing within synchronized cortical layer 2/3 microcircuits therefore appears to robustly signal each active touch response.     

Mesencephalic chronic electrodes in pain patients. An electrophysiological study.

Early components of lemniscal potentials after contralateral median nerve or mechanical stimulus are due to lemniscal pathways, whereas later components, after 70 msec appearing bilaterally and at higher stimulus intensities probably express extralemniscal activity. Evoked potentials in the central gray matter show much smaller amplitudes compared with somatosensory cortical evoked potentials (SSEP). The strongest component is a negative wave after 70--100 msec. Longer conditioning stimulation of the lemniscal system inhibits late components in the median nerve evoked cortical potentials. On the contrary, stimulation of the nonspecific periaqueductal gray matter produces inhibition of early components of cortical SSEP together with facilitation of late components.     

Somatotopy of human hand somatosensory cortex revealed by dipole source analysis of early somatosensory evoked potentials and 3D-NMR tomography

Somatosensory evoked potentials (SEPs) to median nerve and finger stimulation were analyzed by means of spatio-temporal dipole modelling combined with 3D-NMR tomography in 8 normal subjects. The early SEPs were modelled by 3 equivalent dipoles located in the region of the brain-stem (B) and in the region of the contralateral somatosensory cortex (T and R). Dipole B explained peaks P14 and N18 at the scalp. Dipole T was tangentially oriented and explained the N20-P20, dipole R was radially oriented and modelled the P22. The tangential dipole sources T were located within a distance of 6 mm on the average and all were less than 9 mm from the posterior bank of the central sulcus. In 6 subjects the tangential sources related to finger stimulation arranged along the central sulcus according to the known somatotopy. The radial sources did not show a consistent somatotopic alignment across subjects. We conclude that the combination of dipole source analysis and 3D-NMR tomography is a useful tool for functional localization within the human hand somatosensory cortex.