Response planning in word typing: evidence for inhibition.

Typing is a pervasive phenomenon, yet the underlying neural processes have hardly been studied. Here, the mechanisms of keystroke preparation were studied with a typed picture-naming task performed by expert typists. Electroencephalographic activities recorded over sensorimotor areas prior to first-keystroke onset were examined with time-frequency and event-related potential (ERP) analyses. In the time-frequency domain, a beta event-related desynchronization was present bilaterally. In the ERP analyses, the activity was asymmetric, with negativity and positivity patterns developing over, respectively, contra- and ipsilateral recording sites. This pattern is similar to that observed in choice reaction time tasks, and thus can be interpreted as evidence of contralateral motor cortex activation accompanied by inhibition of the ipsilateral motor cortex. These data constitute the first electrophysiological demonstration of inhibitory activity in typing and pave the way to a thorough study of typing.

High frequency gamma activity in the left hippocampus predicts visual object naming performance.

Access to an object's name requires the retrieval of an arbitrary association between it's identity and a word-label. The hippocampus is essential in retrieving arbitrary associations, and thus could be involved in retrieving the link between an object and its name. To test this hypothesis we recorded the iEEG signal from epileptic patients, directly implanted in the hippocampus, while they performed a picture naming task. High-frequency broadband gamma (50-150 Hz) responses were computed as an index of population-level spiking activity. Our results show, for the first time, single-trial hippocampal dynamics between visual confrontation and naming. Remarkably, the latency of the hippocampal response predicts naming latency, while inefficient hippocampal activation is associated with "tip-of-the-tongue" states (a failure to retrieve the name of a recognized object) suggesting that the hippocampus is an active component of the naming network and that its dynamics are closely related to efficient word production.

Selective attention modulates high-frequency activity in the face-processing network.

Face processing depends on the orchestrated activity of a large-scale neuronal network. Its activity can be modulated by attention as a function of task demands. However, it remains largely unknown whether voluntary, endogenous attention and reflexive, exogenous attention to facial expressions equally affect all regions of the face-processing network, and whether such effects primarily modify the strength of the neuronal response, the latency, the duration, or the spectral characteristics. We exploited the good temporal and spatial resolution of intracranial electroencephalography (iEEG) and recorded from depth electrodes to uncover the fast dynamics of emotional face processing. We investigated frequency-specific responses and event-related potentials (ERP) in the ventral occipito-temporal cortex (VOTC), ventral temporal cortex (VTC), anterior insula, orbitofrontal cortex (OFC), and amygdala when facial expressions were task-relevant or task-irrelevant. All investigated regions of interest (ROI) were clearly modulated by task demands and exhibited stronger changes in stimulus-induced gamma band activity (50-150 Hz) when facial expressions were task-relevant. Observed latencies demonstrate that the activation is temporally coordinated across the network, rather than serially proceeding along a processing hierarchy. Early and sustained responses to task-relevant faces in VOTC and VTC corroborate their role for the core system of face processing, but they also occurred in the anterior insula. Strong attentional modulation in the OFC and amygdala (300 msec) suggests that the extended system of the face-processing network is only recruited if the task demands active face processing. Contrary to our expectation, we rarely observed differences between fearful and neutral faces. Our results demonstrate that activity in the face-processing network is susceptible to the deployment of selective attention. Moreover, we show that endogenous attention operates along the whole face-processing network, and that these effects are reflected in frequency-specific changes in the gamma band.

Functional selectivity in the human occipitotemporal cortex during natural vision: evidence from combined intracranial EEG and eye-tracking.

Eye movements are a constant and essential component of natural vision, yet, most of our knowledge about the human visual system comes from experiments that restrict them. This experimental constraint is mostly in place to control visual stimuli presentation and to avoid artifacts in non-invasive measures of brain activity, however, this limitation can be overcome with intracranial EEG (iEEG) recorded from epilepsy patients. Moreover, the high-frequency components of the iEEG signal (between about 50 and 150Hz) can provide a proxy of population-level spiking activity in any cortical area during free-viewing. We combined iEEG with high precision eye-tracking to study fine temporal dynamics and functional specificity in the fusiform face (FFA) and visual word form area (VWFA) while patients inspected natural pictures containing faces and text. We defined the first local measure of visual (electrophysiological) responsiveness adapted to free-viewing in humans: amplitude modulations in the high-frequency activity range (50-150Hz) following fixations (fixation-related high-frequency response). We showed that despite the large size of receptive fields in the ventral occipito-temporal cortex, neural activity during natural vision of realistic cluttered scenes is mostly dependent upon the category of the foveated stimulus - suggesting that category-specificity is preserved during free-viewing and that attention mechanisms might filter out the influence of objects surrounding the fovea.

Evidence for, and predictions from, forward modeling in language production.

Pickering & Garrod (P&G) put forward the interesting idea that language production relies on forward modeling operating at multiple processing levels. The evidence currently available to substantiate this idea mostly concerns sensorimotor processes and not more abstract linguistic levels (e.g., syntax, semantics, phonology). The predictions that follow from the claim seem too general, in their current form, to guide specific empirical tests.

How silent is silent reading? Intracerebral evidence for top-down activation of temporal voice areas during reading.

As you might experience it while reading this sentence, silent reading often involves an imagery speech component: we can hear our own "inner voice" pronouncing words mentally. Recent functional magnetic resonance imaging studies have associated that component with increased metabolic activity in the auditory cortex, including voice-selective areas. It remains to be determined, however, whether this activation arises automatically from early bottom-up visual inputs or whether it depends on late top-down control processes modulated by task demands. To answer this question, we collaborated with four epileptic human patients recorded with intracranial electrodes in the auditory cortex for therapeutic purposes, and measured high-frequency (50-150 Hz) "gamma" activity as a proxy of population level spiking activity. Temporal voice-selective areas (TVAs) were identified with an auditory localizer task and monitored as participants viewed words flashed on screen. We compared neural responses depending on whether words were attended or ignored and found a significant increase of neural activity in response to words, strongly enhanced by attention. In one of the patients, we could record that response at 800 ms in TVAs, but also at 700 ms in the primary auditory cortex and at 300 ms in the ventral occipital temporal cortex. Furthermore, single-trial analysis revealed a considerable jitter between activation peaks in visual and auditory cortices. Altogether, our results demonstrate that the multimodal mental experience of reading is in fact a heterogeneous complex of asynchronous neural responses, and that auditory and visual modalities often process distinct temporal frames of our environment at the same time.

Long-distance amplitude correlations in the high γ band reveal segregation and integration within the reading network.

Reading sentences involves a distributed network of brain regions acting in concert surrounding the left sylvian fissure. The mechanisms of neural communication underlying the extraction and integration of verbal information across subcomponents of this reading network are still largely unknown. We recorded intracranial EEG activity in 12 epileptic human patients performing natural sentence reading and analyzed long-range corticocortical interactions between local neural activations. During a simple task contrasting semantic, phonological, and purely visual processes, we found process-specific neural activity elicited at the single-trial level, characterized by energy increases in a broad gamma band (40-150 Hz). Correlation analysis between task-induced gamma-band activations revealed a selective fragmentation of the network into specialized subnetworks supporting sentence-level semantic analysis and phonological processing. We extend the implications of our results beyond reading, to propose that gamma-band amplitude correlations might constitute a fundamental mechanism for large-scale neural integration during high-level cognition.

Auditory cortex basal activity modulates cochlear responses in chinchillas.

BACKGROUND: The auditory efferent system has unique neuroanatomical pathways that connect the cerebral cortex with sensory receptor cells. Pyramidal neurons located in layers V and VI of the primary auditory cortex constitute descending projections to the thalamus, inferior colliculus, and even directly to the superior olivary complex and to the cochlear nucleus. Efferent pathways are connected to the cochlear receptor by the olivocochlear system, which innervates outer hair cells and auditory nerve fibers. The functional role of the cortico-olivocochlear efferent system remains debated. We hypothesized that auditory cortex basal activity modulates cochlear and auditory-nerve afferent responses through the efferent system. METHODOLOGY/PRINCIPAL FINDINGS: Cochlear microphonics (CM), auditory-nerve compound action potentials (CAP) and auditory cortex evoked potentials (ACEP) were recorded in twenty anesthetized chinchillas, before, during and after auditory cortex deactivation by two methods: lidocaine microinjections or cortical cooling with cryoloops. Auditory cortex deactivation induced a transient reduction in ACEP amplitudes in fifteen animals (deactivation experiments) and a permanent reduction in five chinchillas (lesion experiments). We found significant changes in the amplitude of CM in both types of experiments, being the most common effect a CM decrease found in fifteen animals. Concomitantly to CM amplitude changes, we found CAP increases in seven chinchillas and CAP reductions in thirteen animals. Although ACEP amplitudes were completely recovered after ninety minutes in deactivation experiments, only partial recovery was observed in the magnitudes of cochlear responses. CONCLUSIONS/SIGNIFICANCE: These results show that blocking ongoing auditory cortex activity modulates CM and CAP responses, demonstrating that cortico-olivocochlear circuits regulate auditory nerve and cochlear responses through a basal efferent tone. The diversity of the obtained effects suggests that there are at least two functional pathways from the auditory cortex to the cochlea.

Efficient "pop-out" visual search elicits sustained broadband γ activity in the dorsal attention network.

An object that differs markedly from its surrounding-for example, a red cherry among green leaves-seems to pop out effortlessly in our visual experience. The rapid detection of salient targets, independently of the number of other items in the scene, is thought to be mediated by efficient search brain mechanisms. It is not clear, however, whether efficient search is actually an "effortless" bottom-up process or whether it also involves regions of the prefrontal cortex generally associated with top-down sustained attention. We addressed this question with intracranial EEG (iEEG) recordings designed to identify brain regions underlying a classic visual search task and correlate neural activity with target detection latencies on a trial-by-trial basis with high temporal precision recordings of these regions in epileptic patients. The spatio-temporal dynamics of single-trial spectral analysis of iEEG recordings revealed sustained energy increases in a broad gamma band (50-150 Hz) throughout the duration of the search process in the entire dorsal attention network both in efficient and inefficient search conditions. By contrast to extensive theoretical and experimental indications that efficient search relies exclusively on transient bottom-up processes in visual areas, we found that efficient search is mediated by sustained gamma activity in the dorsal lateral prefrontal cortex and the anterior cingulate cortex, alongside the superior parietal cortex and the frontal eye field. Our findings support the hypothesis that active visual search systematically involves the frontal-parietal attention network and therefore, executive attention resources, regardless of target saliency.

Reading the mind's eye: online detection of visuo-spatial working memory and visual imagery in the inferior temporal lobe.

Several brain regions involved in visual perception have been shown to also participate in non-sensory cognitive processes of visual representations. Here we studied the role of ventral visual pathway areas in visual imagery and working memory. We analyzed intracerebral EEG recordings from the left inferior temporal lobe of an epileptic patient during working memory tasks and mental imagery. We found that high frequency gamma-band activity (50-150 Hz) in the inferior temporal gyrus (ITG) increased with memory load only during visuo-spatial, but not verbal, working memory. Using a real-time set-up to measure and visualize gamma-band activity online--BrainTV--we found a systematic activity increase in ITG when the patient was visualizing a letter (visual imagery), but not during perception of letters. In contrast, only 7 mm more medially, neurons located in the fusiform gyrus exhibited a complete opposite pattern, responding during verbal working memory retention and letter presentation, but not during imagery or visuo-spatial working memory maintenance. Talairach coordinates indicate that the fusiform contact site corresponds to the word form area, suggesting that this region has a role not only in processing letter-strings, but also in working memory retention of verbal information. We conclude that neural networks supporting imagination of a visual element are not necessarily the same as those underlying perception of that element. Additionally, we present evidence that gamma-band activity in the inferior temporal lobe, can be used as a direct measure of the efficiency of top-down attentional control over visual areas with implications for the development of novel brain-computer interfaces. Finally, by just reading gamma-band activity in these two recording sites, it is possible to determine, accurately and in real-time, whether a given memory content is verbal or visuo-spatial.

Neural mechanisms of human perceptual learning: electrophysiological evidence for a two-stage process.

BACKGROUND: Humans and other animals change the way they perceive the world due to experience. This process has been labeled as perceptual learning, and implies that adult nervous systems can adaptively modify the way in which they process sensory stimulation. However, the mechanisms by which the brain modifies this capacity have not been sufficiently analyzed. METHODOLOGY/PRINCIPAL FINDINGS: We studied the neural mechanisms of human perceptual learning by combining electroencephalographic (EEG) recordings of brain activity and the assessment of psychophysical performance during training in a visual search task. All participants improved their perceptual performance as reflected by an increase in sensitivity (d') and a decrease in reaction time. The EEG signal was acquired throughout the entire experiment revealing amplitude increments, specific and unspecific to the trained stimulus, in event-related potential (ERP) components N2pc and P3 respectively. P3 unspecific modification can be related to context or task-based learning, while N2pc may be reflecting a more specific attentional-related boosting of target detection. Moreover, bell and U-shaped profiles of oscillatory brain activity in gamma (30-60 Hz) and alpha (8-14 Hz) frequency bands may suggest the existence of two phases for learning acquisition, which can be understood as distinctive optimization mechanisms in stimulus processing. CONCLUSIONS/SIGNIFICANCE: We conclude that there are reorganizations in several neural processes that contribute differently to perceptual learning in a visual search task. We propose an integrative model of neural activity reorganization, whereby perceptual learning takes place as a two-stage phenomenon including perceptual, attentional and contextual processes.

Task-related gamma-band dynamics from an intracerebral perspective: review and implications for surface EEG and MEG.

Although non-invasive techniques provide functional activation maps at ever-growing spatio-temporal precision, invasive recordings offer a unique opportunity for direct investigations of the fine-scale properties of neural mechanisms in focal neuronal populations. In this review we provide an overview of the field of intracranial Electroencephalography (iEEG) and discuss its strengths and limitations and its relationship to non-invasive brain mapping techniques. We discuss the characteristics of invasive data acquired from implanted epilepsy patients using stereotactic-electroencephalography (SEEG) and electrocorticography (ECoG) and the use of spectral analysis to reveal task-related modulations in multiple frequency components. Increasing evidence suggests that gamma-band activity (>40 Hz) might be a particularly efficient index for functional mapping. Moreover, the detection of high gamma activity may play a crucial role in bridging the gap between electrophysiology and functional imaging studies as well as in linking animal and human data. The present review also describes recent advances in real-time invasive detection of oscillatory modulations (including gamma activity) in humans. Furthermore, the implications of intracerebral findings on future non-invasive studies are discussed.

A neutral cue facilitates detection of a visual target by modulating attention

Twelve rats were trained to perform a two-choice visual detection task in which a right or left light was presented and the animals were required to press the lever located under the illuminated light for a food reward. In seventy percent of the trials the target light was preceded by presentation of a neutral cue (a central light). Relevance of the neutral cue for detection of the target was analyzed by comparing behavioral indices of attention in its presence and absence. Accuracy was significantly higher in presence than in absence of the neutral cue, while mean response latencies were lower in presence than in absence of the neutral cue. These results indicate that the animals allocated attentional resources on the target detection during a high expectancy period after the onset of the neutral cue. This could facilitate target detection and improve the performance in the presence of the neutral cue.

A neutral cue facilitates detection of a visual target by modulating attention.

Twelve rats were trained to perform a two-choice visual detection task in which a right or left light was presented and the animals were required to press the lever located under the illuminated light for a food reward. In seventy percent of the trials the target light was preceded by presentation of a neutral cue (a central light). Relevance of the neutral cue for detection of the target was analyzed by comparing behavioral Indices of attention in its presence and absence. Accuracy was significantly higher in presence than in absence of the neutral cue, while mean response latencies were lower in presence than in absence of the neutral cue. These results indicate that the animals allocated attentional resources on the target detection during a high expectancy period after the onset of the neutral cue. This could facilitate target detection and improve the performance in the presence of the neutral cue.

Selective attention to visual stimuli reduces cochlear sensitivity in chinchillas.

It is generally accepted that during periods of attention to specific stimuli there are changes in the neural activity of central auditory structures; however, it is controversial whether attention can modulate auditory responses at the cochlear level. Several studies performed in animals as well as in humans have attempted to find a modulation of cochlear responses during visual attention with contradictory results. Here, we have appraised cochlear sensitivity in behaving chinchillas by measuring, with a chronically implanted round-window electrode, sound-evoked auditory-nerve compound action potentials and cochlear microphonics, a measure of outer hair cell function, during selective attention to visual stimuli. Chinchillas were trained in a visual discrimination or in an auditory frequency discrimination two-choice task. We found a significant decrease of cochlear sensitivity during the period of attention to visual stimuli in the animals performing the visual discrimination task, but not in those performing the auditory task, demonstrating that this physiological effect is related to selective attention to visual stimuli rather than to an increment in arousal level. Furthermore, the magnitude of the cochlear-sensitivity reductions increased in sessions performed with shorter target-light durations (4-0.5 s), suggesting that this effect is stronger for higher attentional demands of the task. These results demonstrate that afferent auditory activity is modulated by selective attention as early as at sensory transduction, possibly through activation of olivocochlear efferent fibers.

Relevance of a neutral cue in a two-choice detection task in the rat.

Relevance of a neutral cue for performance in a two-choice visuospatial detection task was examined. Nine rats were trained, 5 with short intertrial interval (ITI) and 4 with long ITI, to detect a target (lateral lights) presented after a neutral-cue (central light). The removal of the neutral-cue decreased accuracy and increased response latencies and omissions. These results demonstrate that a neutral-cue, preceding the target, is relevant for the performance, suggesting that rats are highly expectant during the neutral-cue and reallocate attentional resources during ITI. Furthermore, latencies were higher, omissions were lower and the fall of accuracy was greater for rats with long than with short ITI, which could indicate that the neutral-cue was more relevant for the former group.