A real-time marker of object-based attention in the human brain. A possible component of a "gate-keeping mechanism" performing late attentional selection in the Ventro-Lateral Prefrontal Cortex.

The decision to process an incoming stimulus attentively - and to trigger a follow-up cascade of high-level processes - is strategic for the human brain as it becomes transiently unavailable to subsequent stimulus processing. In this study, we set to identify brain networks that carry out such evaluations. We therefore assessed the time-course of neural responses with intracerebral EEG in human patients during an attentional reading task, contrasting to-be-attended vs. to-be-ignored items. We measured High-Frequency Activity [50-150 â€‹Hz] as a proxy of population-level spiking activity and we identified a crucial component of a Gate-Keeping Mechanism bilateral in the mid-Ventro-Lateral Prefrontal Cortex (VLPFC), at the interplay of the Ventral and Dorsal Attention Networks, that selectively reacts before domain specialized cortical regions that engage in full stimulus analysis according to task demands.

What is that little voice inside my head? Inner speech phenomenology, its role in cognitive performance, and its relation to self-monitoring.

The little voice inside our head, or inner speech, is a common everyday experience. It plays a central role in human consciousness at the interplay of language and thought. An impressive host of research works has been carried out on inner speech these last fifty years. Here we first describe the phenomenology of inner speech by examining five issues: common behavioural and cerebral correlates with overt speech, different types of inner speech (wilful verbal thought generation and verbal mind wandering), presence of inner speech in reading and in writing, inner signing and voice-hallucinations in deaf people. Secondly, we review the role of inner speech in cognitive performance (i.e., enhancement vs. perturbation). Finally, we consider agency in inner speech and how our inner voice is known to be self-generated and not produced by someone else.

Turning visual shapes into sounds: early stages of reading acquisition revealed in the ventral occipitotemporal cortex.

The exact role of the left ventral occipitotemporal cortex (VOTC) during the initial stages of reading acquisition is a hotly debated issue, especially regarding the comparative effect of learning on early stimulus-dependent vs. later task-dependent processes. We show that this controversy can be solved with high-temporal resolution intracerebral EEG recordings of the VOTC. We measured High-Frequency Activity (50-150 Hz) as a proxy of population-level spiking activity while participants learned Japanese Katakana symbols, and found that learning primarily affects top-down/task-dependent neural processing, after a few minutes only. In contrast, adaptation of early bottom-up/stimulus-dependent processing takes several days to adapt and provides the basis for fluent reading. Such evidence that two consecutive stages of neural processing, stimulus- and task-dependent are differentially affected by learning, can reconcile seemingly opposite hypotheses on the role of the VOTC during reading acquisition.

Silence is golden: transient neural deactivation in the prefrontal cortex during attentive reading.

It is becoming increasingly clear that attention-demanding tasks engage not only activation of specific cortical regions but also deactivation of other regions that could interfere with the task at hand. At the same time, electrophysiological studies in animals and humans have found that the participation of cortical regions to cognitive processes translates into local synchronization of rhythmic neural activity at frequencies above 40 Hz (so-called gamma-band synchronization). Such synchronization is seen as a potential facilitator of neural communication and synaptic plasticity. We found evidence that cognitive processes can also involve the disruption of gamma-band activity in high-order brain regions. Intracerebral electroencephalograms were recorded in 3 epileptic patients during 2 reading tasks. Visual presentation of words induced a strong deactivation in a broad (20-150 Hz) frequency range in the left ventral lateral prefrontal cortex, in parallel with gamma-band activations within the reading network, including Broca's area. The observed energy decrease in neural signals was reproducible across patients. It peaked around 500 ms after stimulus onset and appeared subject to attention-modulated amplification. Our results suggest that cognition might be mediated by a coordinated interaction between regional gamma-band synchronizations and desynchronizations, possibly reflecting enhanced versus reduced local neural communication.

From autopoiesis to neurophenomenology: Francisco Varela's exploration of the biophysics of being

This paper reviews in detail Francisco Varela's work on subjectivity and consciousness in the biological sciences. His original approach to this [quot ]hard problem[quot ] presents a subjectivity that is radically intertwined with its biological and physical roots. It must be understood within the framework of his theory of a concrete, embodied dynamics, grounded in his general theory of autonomous systems. Through concepts and paradigms such as biological autonomy, embodiment and neurophenomenology, the article explores the multiple levels of circular causality assumed by Varela to play a fundamental role in the emergence of human experience. The concept of biological autonomy provides the necessary and sufficient conditions for characterizing biological life and identity as an emergent and circular self-producing process. Embodiment provides a systemic and dynamical framework for understanding how a cognitive self--a mind--can arise in an organism in the midst of its operational cycles of internal regulation and ongoing sensorimotor coupling. Global subjective properties can emerge at different levels from the interactions of components and can reciprocally constrain local processes through an ongoing, recursive morphodynamics. Neurophenomenology is a supplementary step in the study of consciousness. Through a rigorous method, it advocates the careful examination of experience with first-person methodologies. It attempts to create heuristic mutual constraints between biophysical data and data produced by accounts of subjective experience. The aim is to explicitly ground the active and disciplined insight the subject has about his/her experience in a biophysical emergent process. Finally, we discuss Varela's essential contribution to our understanding of the generation of consciousness in the framework of what we call his "biophysics of being".

The brainweb: phase synchronization and large-scale integration.

The emergence of a unified cognitive moment relies on the coordination of scattered mosaics of functionally specialized brain regions. Here we review the mechanisms of large-scale integration that counterbalance the distributed anatomical and functional organization of brain activity to enable the emergence of coherent behaviour and cognition. Although the mechanisms involved in large-scale integration are still largely unknown, we argue that the most plausible candidate is the formation of dynamic links mediated by synchrony over multiple frequency bands.

A quantitative study of gamma-band activity in human intracranial recordings triggered by visual stimuli.

This paper studies gamma-band responses from two implanted epileptic patients during a simple visual discrimination task. Our main aim was to ascertain, in a reliable manner, whether evoked (stimulus-locked) and induced (triggered by, but not locked to, stimuli) responses are present in intracranial recordings. For this purpose, we introduce new methods adapted to detect the presence of gamma responses at this level of recording, intermediary between EEG-scalp and unicellular responses. The analysis relies on a trial-by-trial time-frequency analysis and on the use of surrogate data for statistical testing. We report that visual stimulation reliably elicits evoked and induced responses in human intracranial recordings. Induced intracranial gamma activity is significantly present in short oscillatory bursts (a few cycles) following visual stimulation. These responses are highly variable from trial to trial, beginning after 200 ms and lasting up to 500 ms. In contrast, intracranial-evoked gamma responses concentrate around 100 ms latencies corresponding to evoked responses observed on the scalp. We discuss our results in relation to scalp gamma response in a similar protocol [Tallon-Baudry et al. (1996) J. Neurosci., 16, 4240-4249] and draw some conclusions for bridging the gap between gamma oscillations observed on the scalp surface and their possible cortical sources.

Inter- and intralimb oscillator coupling in parkinsonian tremor.

This study reports the findings of an analysis of temporal correlation between tremor of different muscles of the same and different limbs in four patients with Parkinson's disease. Spectral coherence methods were used for determining whether simultaneously occurring oscillations in the electromyograms of different muscles are statistically coupled. The incidence of significant coherence was considerably higher for muscle pairs in the same limb than for pairs in different limbs; Parkinson's disease tremor is coupled within but not between limbs. Because the characteristics of tremor are known to vary under different behavioral situations, the intralimb coupling was examined for different tasks. A mental arithmetic task resulted in an increase in the coherence between muscles of the same limb, whereas the finger-to-nose task decreased the coherence. No significant change in coherence was found for a postural task. The amplitude and regularity of tremor electromyography showed changes analogous to those in coherence. These results support the hypothesis that tremor in different limbs results from the activity of several neural circuits oscillating independently. The results also emphasize the value of these methods for rigorously characterizing tremor, in relation to disease state, behavioral conditions, and the selection of treatment strategies.

Why bother to spatially embed EEG? Comments on Pritchard et al., Psychophysiology, 33, 362-368, 1996.

In a recent paper, Pritchard, Krieble, and Duke (Psychophysiology, 33, 362-368, 1996) studied the validity of spatial embedding of electroencephalographic (EEG) data and rejected this method in favor of time-delay embedding. The present paper describes the nonlinear characterization of brain dynamics using either spatial or time-delay embedding. We discuss the arguments published in Pritchard et al. (1996) and demonstrate that the spatial embedding cannot be rejected on this basis. We also point out the limitations of both spatial and time-delay embeddings related to the spatial extension and the high-dimensional dynamics of brain activity.

Perception's shadow: long-distance synchronization of human brain activity.

Transient periods of synchronization of oscillating neuronal discharges in the frequency range 30-80 Hz (gamma oscillations) have been proposed to act as an integrative mechanism that may bring a widely distributed set of neurons together into a coherent ensemble that underlies a cognitive act. Results of several experiments in animals provide support for this idea. In humans, gamma oscillations have been described both on the scalp (measured by electroencephalography and magnetoencephalography) and in intracortical recordings, but no direct participation of synchrony in a cognitive task has been demonstrated so far. Here we record electrical brain activity from subjects who are viewing ambiguous visual stimuli (perceived either as faces or as meaningless shapes). We show for the first time, to our knowledge, that only face perception induces a long-distance pattern of synchronization, corresponding to the moment of perception itself and to the ensuing motor response. A period of strong desynchronization marks the transition between the moment of perception and the motor response. We suggest that this desynchronization reflects a process of active uncoupling of the underlying neural ensembles that is necessary to proceed from one cognitive state to another.

Measuring phase synchrony in brain signals.

This article presents, for the first time, a practical method for the direct quantification of frequency-specific synchronization (i.e., transient phase-locking) between two neuroelectric signals. The motivation for its development is to be able to examine the role of neural synchronies as a putative mechanism for long-range neural integration during cognitive tasks. The method, called phase-locking statistics (PLS), measures the significance of the phase covariance between two signals with a reasonable time-resolution (<100 ms). Unlike the more traditional method of spectral coherence, PLS separates the phase and amplitude components and can be directly interpreted in the framework of neural integration. To validate synchrony values against background fluctuations, PLS uses surrogate data and thus makes no a priori assumptions on the nature of the experimental data. We also apply PLS to investigate intracortical recordings from an epileptic patient performing a visual discrimination task. We find large-scale synchronies in the gamma band (45 Hz), e.g., between hippocampus and frontal gyrus, and local synchronies, within a limbic region, a few cm apart. We argue that whereas long-scale effects do reflect cognitive processing, short-scale synchronies are likely to be due to volume conduction. We discuss ways to separate such conduction effects from true signal synchrony.

Temporal patterns in human epileptic activity are modulated by perceptual discriminations.

We studied subdural recordings from a patient with an unusually focal and stable occipito-temporal epileptic discharge under four experimental conditions. The series of time intervals between successive spike discharges displayed a few (3-5) clusters of periodic values representing statistically significant short-term periodicities when tested against surrogate data. This short-term predictability was modulated during the different experimental conditions by periodicity shifts of the order of 15-30 ms. Correspondingly, there was an increased gamma-band (30-70 Hz) coherence between the epileptic focus and surrounding recording sites. We conclude that the focal epileptic activity is part of an extended network of neural activities which exert a fast modulation reflected in changes of transiently periodic activities.

Spatial extension of brain activity fools the single-channel reconstruction of EEG dynamics.

We report here on a first attempt to settle the methodological controversy between advocates of two alternative reconstruction approaches for temporal dynamics in brain signals: the single-channel method (using data from one recording site and reconstructing by time-lags), and the multiple-channel method (using data from a spatially distributed set of recordings sites and reconstructing by means of spatial position). For the purpose of a proper comparison of these two techniques, we computed a series of EEG-like measures on the basis of well-known dynamical systems placed inside a spherical model of the head. For each of the simulations, the correlation dimension estimates obtained by both methods were calculated and compared, when possible, with the known (or estimated) dimension of the underlying dynamical system. We show that the single-channel method fails to reliably quantify spatially extended dynamics, while the multichannel method performs better. It follows that the latter is preferable, given the known spatially distributed nature of brain processes. Hum. Brain Mapping 5:26-47, 1997. (c) 1997 Wiley-Liss, Inc.