Nelson J. Trujillo-Barreto

Bayesian model averaging in EEG/MEG imaging.

In this paper, the Bayesian Theory is used to formulate the Inverse Problem (IP) of the EEG/MEG. This formulation offers a comparison framework for the wide range of inverse methods available and allows us to address the problem of model uncertainty that arises when dealing with different solutions for a single data. In this case, each model is defined by the set of assumptions of the inverse method used, as well as by the functional dependence between the data and the Primary Current Density (PCD) inside the brain. The key point is that the Bayesian Theory not only provides for posterior estimates of the parameters of interest (the PCD) for a given model, but also gives the possibility of finding posterior expected utilities unconditional on the models assumed. In the present work, this is achieved by considering a third level of inference that has been systematically omitted by previous Bayesian formulations of the IP. This level is known as Bayesian model averaging (BMA). The new approach is illustrated in the case of considering different anatomical constraints for solving the IP of the EEG in the frequency domain. This methodology allows us to address two of the main problems that affect linear inverse solutions (LIS): (a) the existence of ghost sources and (b) the tendency to underestimate deep activity. Both simulated and real experimental data are used to demonstrate the capabilities of the BMA approach, and some of the results are compared with the solutions obtained using the popular low-resolution electromagnetic tomography (LORETA) and its anatomically constraint version (cLORETA).

Directed Cortical Information Flow during Human Object Recognition: Analyzing Induced EEG Gamma-Band Responses in Brain's Source Space

The increase of induced gamma-band responses (iGBRs; oscillations >30 Hz) elicited by familiar (meaningful) objects is well established in electroencephalogram (EEG) research. This frequency-specific change at distinct locations is thought to indicate the dynamic formation of local neuronal assemblies during the activation of cortical object representations. As analytically power increase is just a property of a single location, phase-synchrony was introduced to investigate the formation of large-scale networks between spatially distant brain sites. However, classical phase-synchrony reveals symmetric, pair-wise correlations and is not suited to uncover the directionality of interactions. Here, we investigated the neural mechanism of visual object processing by means of directional coupling analysis going beyond recording sites, but rather assessing the directionality of oscillatory interactions between brain areas directly. This study is the first to identify the directionality of oscillatory brain interactions in source space during human object recognition and suggests that familiar, but not unfamiliar, objects engage widespread reciprocal information flow. Directionality of cortical information-flow was calculated based upon an established Granger-Causality coupling-measure (partial-directed coherence; PDC) using autoregressive modeling. To enable comparison with previous coupling studies lacking directional information, phase-locking analysis was applied, using wavelet-based signal decompositions. Both, autoregressive modeling and wavelet analysis, revealed an augmentation of iGBRs during the presentation of familiar objects relative to unfamiliar controls, which was localized to inferior-temporal, superior-parietal and frontal brain areas by means of distributed source reconstruction. The multivariate analysis of PDC evaluated each possible direction of brain interaction and revealed widespread reciprocal information-transfer during familiar object processing. In contrast, unfamiliar objects entailed a sparse number of only unidirectional connections converging to parietal areas. Considering the directionality of brain interactions, the current results might indicate that successful activation of object representations is realized through reciprocal (feed-forward and feed-backward) information-transfer of oscillatory connections between distant, functionally specific brain areas.

Realistically Coupled Neural Mass Models Can Generate EEG Rhythms

We study the generation of EEG rhythms by means of realistically coupled neural mass models. Previous neural mass models were used to model cortical voxels and the thalamus. Interactions between voxels of the same and other cortical areas and with the thalamus were taken into account. Voxels within the same cortical area were coupled (short-range connections) with both excitatory and inhibitory connections, while coupling between areas (long-range connections) was considered to be excitatory only. Short-range connection strengths were modeled by using a connectivity function depending on the distance between voxels. Coupling strength parameters between areas were defined from empirical anatomical data employing the information obtained from probabilistic paths, which were tracked by water diffusion imaging techniques and used to quantify white matter tracts in the brain. Each cortical voxel was then described by a set of 16 random differential equations, while the thalamus was described by a set of 12 random differential equations. Thus, for analyzing the neuronal dynamics emerging from the interaction of several areas, a large system of differential equations needs to be solved. The sparseness of the estimated anatomical connectivity matrix reduces the number of connection parameters substantially, making the solution of this system faster. Simulations of human brain rhythms were carried out in order to test the model. Physiologically plausible results were obtained based on this anatomically constrained neural mass model.

Biophysical model for integrating neuronal activity, EEG, fMRI and metabolism

Our goal is to model the coupling between neuronal activity, cerebral metabolic rates of glucose and oxygen consumption, cerebral blood flow (CBF), electroencephalography (EEG) and blood oxygenation level-dependent (BOLD) responses. In order to accomplish this, two previous models are coupled: a metabolic/hemodynamic model (MHM) for a voxel, linking BOLD signals and neuronal activity, and a neural mass model describing the neuronal dynamics within a voxel and its interactions with voxels of the same area (short-range interactions) and other areas (long-range interactions). For coupling both models, we take as the input to the BOLD model, the number of active synapses within the voxel, that is, the average number of synapses that will receive an action potential within the time unit. This is obtained by considering the action potentials transmitted between neuronal populations within the voxel, as well as those arriving from other voxels. Simulations are carried out for testing the integrated model. Results show that realistic evoked potentials (EP) at electrodes on the scalp surface and the corresponding BOLD signals for each voxel are produced by the model. In another simulation, the alpha rhythm was reproduced and reasonable similarities with experimental data were obtained when calculating correlations between BOLD signals and the alpha power curve. The origin of negative BOLD responses and the characteristics of EEG, PET and BOLD signals in Alzheimers disease were also studied.

Bayesian M/EEG source reconstruction with spatio-temporal priors

This article proposes a Bayesian spatio-temporal model for source reconstruction of M/EEG data. The usual two-level probabilistic model implicit in most distributed source solutions is extended by adding a third level which describes the temporal evolution of neuronal current sources using time-domain General Linear Models (GLMs). These comprise a set of temporal basis functions which are used to describe event-related M/EEG responses. This places M/EEG analysis in a statistical framework that is very similar to that used for PET and fMRI. The experimental design can be coded in a design matrix, effects of interest characterized using contrasts and inferences made using posterior probability maps. Importantly, as is the case for single-subject fMRI analysis, trials are treated as fixed effects and the approach takes into account between-trial variance, allowing valid inferences to be made on single-subject data. The proposed probabilistic model is efficiently inverted by using the Variational Bayes framework under a convenient mean-field approximation (VB-GLM). The new method is tested with biophysically realistic simulated data and the results are compared to those obtained with traditional spatial approaches like the popular Low Resolution Electromagnetic TomogrAphy (LORETA) and minimum variance Beamformer. Finally, the VB-GLM approach is used to analyze an EEG data set from a face processing experiment.

Brain electrical tomography (BET) analysis of induced gamma band responses during a simple object recognition task

The formation of cortical object representations requires the activation of cell assemblies, correlated by induced oscillatory bursts above 20 Hz (gamma band), which are characterized by trial-by-trial latency fluctuations around a mean of approximately 300 ms after stimulus onset. The present electroencephalogram (EEG) study was intended to uncover to the generators of induced gamma band responses (GBRs) and to analyze phase-synchronization between these sources. A standard object recognition task was used to elicit gamma activity. At the scalp surface (electrode space), we found an augmentation of induced GBRs after the presentation of meaningful (familiar) as opposed to meaningless (unfamiliar) stimuli, which was accompanied by a dense pattern of significant phase-locking values between distant recording sites. Subsequently, intracranial current density distributions compatible with the observed scalp voltage topographies were estimated by means of VARETA (Variable Resolution Electromagnetic Tomography). In source space brain electrical tomographies (BETs) revealed widespread generators of induced GBRs at temporal, parietal, posterior, and frontal areas. Phase-locking analysis was calculated between re-constructed electrode signals based on separate forward solutions of the observed generators, thereby eliminating the possibly confounding influence of activity from areas not under observation. The results support the view that induced GBRs signify synchronous neuronal activity in a broadly distributed network during object recognition. The localization of the generators of event-related potentials (ERPs), evoked gamma activity, and induced alpha activity revealed different sources as compared to the induced GBR and, thus, seem to mirror complementary functions during the present task as compared to induced high-frequency brain dynamics.

Rapid extraction of auditory feature contingencies

Contingent relations between sensory events render the environment predictable and thus facilitate adaptive behavior. The human capacity to detect such relations has been comprehensively demonstrated in paradigms in which contingency rules were task-relevant or in which they applied to motor behavior. The extent to which contingencies can also be extracted from events that are unrelated to the current goals of the organism has remained largely unclear. The present study addressed the emergence of contingency-related effects for behaviorally irrelevant auditory stimuli and the cortical areas involved in the processing of such contingency rules. Contingent relations between different features of temporally separate events were embedded in a new dynamic protocol. Participants were presented with the auditory stimulus sequences while their attention was captured by a video. The mismatch negativity (MMN) component of the event-related brain potential (ERP) was employed as an electrophysiological correlate of contingency detection. MMN generators were localized by means of scalp current density (SCD) and primary current density (PCD) analyses with variable resolution electromagnetic tomography (VARETA). Results show that task-irrelevant contingencies can be extracted from about fifteen to twenty successive events conforming to the contingent relation. Topographic and tomographic analyses reveal the involvement of the auditory cortex in the processing of contingency violations. The present data provide evidence for the rapid encoding of complex extrapolative relations in sensory areas. This capacity is of fundamental importance for the organism in its attempt to model the sensory environment outside the focus of attention.

Processing of abstract rule violations in audition

The ability to encode rules and to detect rule-violating events outside the focus of attention is vital for adaptive behavior. Our brain recordings reveal that violations of abstract auditory rules are processed even when the sounds are unattended. When subjects performed a task related to the sounds but not to the rule, rule violations impaired task performance and activated a network involving supratemporal, parietal and frontal areas although none of the subjects acquired explicit knowledge of the rule or became aware of rule violations. When subjects tried to behaviorally detect rule violations, the brains automatic violation detection facilitated intentional detection. This shows the brains capacity for abstraction – an important cognitive function necessary to model the world. Our study provides the first evidence for the task-independence (i.e. automaticity) of this ability to encode abstract rules and for its immediate consequences for subsequent mental processes.

Sustained spatial attention to vibration is mediated in primary somatosensory cortex.

Focusing attention to a specific body location has been shown to improve processing of events presented at this body location. One important debate concerns the stage in the somatosensory pathway at which the neural response is modulated when one attends to a tactile stimulus. Previous studies focused on components of the somatosensory evoked potential to transient stimuli, and demonstrated an early cortical attentional modulation. The neural basis of sustained spatial stimulus processing with continuous stimulation remains, however, largely unexplored. A way to approach this topic is to present vibrating stimuli with different frequencies for several seconds simultaneously to different body locations while subjects have to attend to the one or the other location. The amplitude of the somatosensory steady-state evoked potential (SSSEP) elicited by these vibrating stimuli increases with attention. On the basis of 128 electrode recordings, we investigated the topographical distribution and the underlying cortical sources by means of a VARETA approach of this attentional amplitude modulation of the SSSEP. Sustained spatial attention was found to be mediated in primary somatosensory cortex with no differences in SSSEP amplitude topographies between attended and unattended body locations. These result patterns were seen as evidence for a low-level sensory gain control mechanism in tactile spatial attention.

Hearing Silences: Human Auditory Processing Relies on Preactivation of Sound-Specific Brain Activity Patterns

The remarkable capabilities displayed by humans in making sense of an overwhelming amount of sensory information cannot be explained easily if perception is viewed as a passive process. Current theoretical and computational models assume that to achieve meaningful and coherent perception, the human brain must anticipate upcoming stimulation. But how are upcoming stimuli predicted in the brain? We unmasked the neural representation of a prediction by omitting the predicted sensory input. Electrophysiological brain signals showed that when a clear prediction can be formulated, the brain activates a template of its response to the predicted stimulus before it arrives to our senses.