Alpha band (8-13Hz) electroencephalography (EEG) oscillations play a key role in cognition, but their generating mechanisms are still poorly understood. Most studies investigating laminar origins of alpha oscillations have been conducted on animals using invasive intracranial recordings. To relate these findings to human alpha generation, non-invasive techniques need to be developed. Layer functional Magnetic Resonance Imaging (fMRI) at ultra-high field (UHF, 7T) allows for the interrogation of brain responses across cortical depths and combined with simultaneous EEG, provides the opportunity to gain new insight into human alpha generation mechanisms. This work establishes a framework to study the generating mechanisms of electrophysiological signals non-invasively in humans using simultaneous EEG layer-fMRI. Data were acquired on 10 participants during an eyes closed/eyes open paradigm. We showed that in 9/10 participants the quality of EEG and Blood Oxygenation Level Dependent (BOLD) fMRI data were sufficient to observe a significant negative correlation between EEG alpha power and the BOLD signal in visual cortex grey matter to the eyes open/eyes closed task. "Deveining" was performed to overcome the increase in BOLD signal toward the pial surface due to draining veins, and the effects of each of the steps in the deveining analysis on the cortical depth profiles of the negative alpha-BOLD correlations studied. The largest effect was dependent on the exclusion of voxels in the tissue immediately surrounding veins. Following deveining, the cortical depth profiles showed the negative alpha-BOLD correlations were significantly weaker in the middle depths compared with deep and superficial depths. When a boxcar rather than EEG alpha power was used to model the task, this depth-dependence was not seen, suggesting this was specific to spontaneous alpha-power modulations. In conclusion, we have established a method to non-invasively interrogate the origins of electrophysiological signals. Our alpha-BOLD depth profiles suggest the alpha signal to an eyes open-closed task is generated in superficial and deep layers suggesting top-down processes.
BACKGROUND:Prior work has shown that transcranial alternating current stimulation (tACS) of parietooccipital alpha oscillations (8-14 Hz) can modulate working memory (WM) performance as a function of the phase lag to endogenous oscillations. However, leveraging this effect using real-time phase-tuned tACS has not been feasible so far due to stimulation artifacts preventing continuous phase tracking. OBJECTIVES AND HYPOTHESIS:We aimed to develop a system that tracks and adapts the phase lag between tACS and ongoing parietooccipital alpha oscillations in real-time. We hypothesized that such real-time phase-tuned tACS enhances working memory performance, depending on the phase lag. METHODS:We developed real-time phase-tuned closed-loop amplitude-modulated tACS (CLAM-tACS) targeting parietooccipital alpha oscillations. CLAM-tACS was applied at six different phase lags relative to ongoing alpha oscillations while participants (N = 21) performed a working memory task. To exclude that behavioral effects of CLAM-tACS were mediated by other factors such as sensory co-stimulation, a second group of participants (N = 25) received equivalent stimulation of the forehead. RESULTS:WM accuracy improved in a phase lag dependent manner (p = 0.0350) in the group receiving parietooccipital stimulation, with the strongest enhancement observed at 330° phase lag between tACS and ongoing alpha oscillations (p = 0.00273, d = 0.976). Moreover, across participants, modulation of frontoparietal alpha oscillations correlated both in amplitude (p = 0.0248) and phase (p = 0.0270) with the modulation of WM accuracy. No such effects were observed in the control group receiving frontal stimulation. CONCLUSIONS:Our results demonstrate the feasibility and efficacy of real-time phase-tuned CLAM-tACS in modulating both brain activity and behavior, thereby paving the way for further investigation into brain-behavior relationships and the exploration of innovative therapeutic applications.
Transcranial alternating current stimulation (tACS) can influence human perception and behavior, with recent evidence also suggesting its potential impact in clinical settings, but the underlying mechanisms are poorly understood. Behavioral and indirect physiological evidence indicates that phase-dependent constructive and destructive interference between the tACS electric field and ongoing brain oscillations may play an important role, but direct in-vivo validation was infeasible because stimulation artifacts impeded such assessment. Using stimulation artifact source separation (SASS), a real-time compatible artifact suppression approach, we overcame this limitation and provide direct evidence for millisecond-by-millisecond phase-dependent enhancement and suppression of ongoing brain oscillations during amplitude-modulated tACS (AM-tACS) across 29 healthy human volunteers. We found that AM-tACS enhanced and suppressed targeted brain oscillations by 11.7 ± 5.14% and 10.1 ± 4.07% respectively. Millisecond-precise modulation of oscillations predicted modulation of behavior (r = 0.65, p < 0.001). These results not only provide direct evidence for constructive and destructive interference as a key mechanism of AM-tACS but suggest superiority of phase-locked (closed-loop) AM-tACS over conventional (open-loop) AM-tACS to purposefully enhance or suppress brain oscillations. Significance The presented data provide direct evidence for a key mechanism underlying neurophysiological and behavioral effects of transcranial alternating current stimulation (tACS), a broadly used neuromodulation approach that yields promising clinical results but also raised controversies because of its variable effects. Our findings not only elucidate the underlying mechanisms of tACS, but also provide the rationale for closed-loop tACS protocols that will enable targeted enhancement and suppression of brain oscillations related to various brain functions such perception, memory or cognition. Towards this end, we introduce the technical prerequisites to establish millisecond-to-millisecond precise closed-loop tACS protocols that will be important to advance tACS as a neuroscientific and clinical tool, for example in the treatment of neuropsychiatric disorders.
EEG alpha (8-13Hz) oscillations occur throughout the cortex but the generating mechanisms are poorly understood. Opinion is divided between alpha being driven by bottom-up, top-down or both of these processes. Using simultaneous 7T-fMRI-EEG with an eyes open/closed paradigm, we assess the generator of alpha by performing layer-fMRI analysis of GE-BOLD data to determine the strongest BOLD-alpha negative layer correlations. We show that, after accounting for draining vein effects using spatial deconvolution, alpha-BOLD correlations are strongest in the superficial and deep layers suggesting they are predominately driven by top-down processes.
Language comprehension relies on integrating words into progressively more complex structures, like phrases and sentences. This hierarchical structure-building is reflected in rhythmic neural activity across multiple timescales in E/MEG in healthy, awake participants. However, recent studies have shown evidence for this "cortical tracking" of higher-level linguistic structures also in a proportion of unresponsive patients. What does this tell us about these patients' residual levels of cognition and consciousness? Must the listener direct their attention toward higher level speech structures to exhibit cortical tracking, and would selective attention across levels of the hierarchy influence the expression of these rhythms? We investigated these questions in an EEG study of 72 healthy human volunteers listening to streams of monosyllabic isochronous English words that were either unrelated (scrambled condition) or composed of four-word-sequences building meaningful sentences (sentential condition). Importantly, there were no physical cues between four-word-sentences. Rather, boundaries were marked by syntactic structure and thematic role assignment. Participants were divided into three attention groups: from passive listening (passive group) to attending to individual words (word group) or sentences (sentence group). The passive and word groups were initially naïve to the sentential stimulus structure, while the sentence group was not. We found significant tracking at word- and sentence rate across all three groups, with sentence tracking linked to left middle temporal gyrus and right superior temporal gyrus. Goal-directed attention to words did not enhance word-rate-tracking, suggesting that word tracking here reflects largely automatic mechanisms, as was shown for tracking at the syllable-rate before. Importantly, goal-directed attention to sentences relative to words significantly increased sentence-rate-tracking over left inferior frontal gyrus. This attentional modulation of rhythmic EEG activity at the sentential rate highlights the role of attention in integrating individual words into complex linguistic structures. Nevertheless, given the presence of high-level cortical tracking under conditions of lower attentional effort, our findings underline the suitability of the paradigm in its clinical application in patients after brain injury. The neural dissociation between passive tracking of sentences and directed attention to sentences provides a potential means to further characterise the cognitive state of each unresponsive patient.
Accurate early prognostication is vital for appropriate long-term care decisions after traumatic brain injury. While measures of resting-state EEG oscillations and their network properties, derived from graph theory, have been shown to provide clinically useful information regarding diagnosis and recovery in patients with chronic disorders of consciousness, little is known about the value of these network measures when calculated from a standard clinical low-density EEG in the acute phase post-injury. To investigate this link, we first validated a set of measures of oscillatory network features between high-density and low-density resting-state EEG in healthy individuals, thus ensuring accurate estimation of underlying cortical function in clinical recordings from patients. Next, we investigated the relationship between these features and the clinical picture and outcome of a group of 18 patients in acute post-traumatic unresponsive states who were not following commands 2 days+ after sedation hold. While the complexity of the alpha network, as indexed by the standard deviation of the participation coefficients, was significantly related to the patients’ clinical picture at the time of EEG, no network features were significantly related to outcome at 3 or 6 months post-injury. Rather, mean relative alpha power across all electrodes improved the accuracy of outcome prediction at 3 months relative to clinical features alone. These results highlight the link between the alpha rhythm and clinical signs of consciousness and suggest the potential for simple measures of resting-state EEG band power to provide a coarse snapshot of brain health for stratification of patients for rehabilitation, therapy and assessments of both covert and overt cognition.
Objective Patients with traumatic brain injury who fail to obey commands after sedation‐washout pose one of the most significant challenges for neurological prognostication. Reducing prognostic uncertainty will lead to more appropriate care decisions and ensure provision of limited rehabilitation resources to those most likely to benefit. Bedside markers of covert residual cognition, including speech comprehension, may reduce this uncertainty. Methods We recruited 28 patients with acute traumatic brain injury who were 2 to 7 days sedation‐free and failed to obey commands. Patients heard streams of isochronous monosyllabic words that built meaningful phrases and sentences while their brain activity via electroencephalography (EEG) was recorded. In healthy individuals, EEG activity only synchronizes with the rhythm of phrases and sentences when listeners consciously comprehend the speech. This approach therefore provides a measure of residual speech comprehension in unresponsive patients. Results Seventeen and 16 patients were available for assessment with the Glasgow Outcome Scale Extended (GOSE) at 3 months and 6 months, respectively. Outcome significantly correlated with the strength of patients’ acute cortical tracking of phrases and sentences ( r > 0.6, p < 0.007), quantified by inter‐trial phase coherence. Linear regressions revealed that the strength of this comprehension response (beta = 0.603, p = 0.006) significantly improved the accuracy of prognoses relative to clinical characteristics alone (eg, Glasgow Coma Scale [GCS], computed tomography [CT] grade). Interpretation A simple, passive, auditory EEG protocol improves prognostic accuracy in a critical period of clinical decision making. Unlike other approaches to probing covert cognition for prognostication, this approach is entirely passive and therefore less susceptible to cognitive deficits, increasing the number of patients who may benefit. ANN NEUROL 2021;89:646–656
ABSTRACT Language comprehension relies on integrating words into progressively more complex structures, like phrases and sentences. This hierarchical structure building is reflected in rhythmic neural activity across multiple timescales in E/MEG (Ding et al., 2016, 2017). How does selective attention across levels of the hierarchy influence the expression of these rhythms? We investigated these questions in an EEG study of 72 healthy human volunteers listening to streams of monosyllabic isochronous English words that were either unrelated (scrambled condition) or composed of four-word-sequences building meaningful sentences (sentential condition). Importantly, there were no physical cues between four-word-sentences but boundaries were marked by syntactic structure and thematic role assignment. Participants were divided into three attention groups: from passive listening (passive group) to attending to individual words (word group) or sentences (sentence group). The passive and word group were naïve to the sentential structure of the stimulus material, while the sentence group were not. We found significant entrainment at word- and sentence rate across all three groups, with sentence entrainment linked to left middle temporal gyrus and right superior temporal gyrus. Goal-directed attention to words did not enhance word-rate-entrainment suggesting that word entrainment relies on largely automatic mechanisms. Importantly, goal-directed attention to sentences relative to words significantly increased sentence-rate-entrainment over left inferior frontal gyrus. This attentional modulation of rhythmic EEG activity at the sentential level highlights the role of attention in integrating individual words into complex linguistic structures. SIGNIFICANCE STATEMENT Neural activity is known to entrain to physical characteristics of auditory stimuli. However, entrainment also occurs with structures lacking physical cues but rather require comprehension of the stimulus’ meaning – for example, entrainment to sentences in speech even without acoustic gaps separating these higher linguistic structures. We investigated how goal-directed attention to low-level (words) and high-level (sentences) linguistic structures influences entrainment strength. Whilst sentence entrainment occurred independently of selective attention, it increased with goal-directed attention towards sentences. Conversely, no such attentional effect was found for word entrainment. While goal-directed attention towards sentences strengthens entrainment, it is no prerequisite for it to occur, suggesting that low attentional effort is required for sentence comprehension, potentially reflecting the importance of speech in humans.
Perception is facilitated by a hierarchy of expectations generated from context and prior knowledge. In auditory processing, violations of local (within-trial) expectations elicit a mismatch negativity, while violations of global (across-trial) expectations elicit a later positive component (P300). This result is taken as evidence of prediction errors ascending through the expectation hierarchy. However, in language comprehension, there is no evidence that violations of semantic expectations across local-global levels similarly elicit a sequence of hierarchical error signals – thus drawing into question the putative link between event-related potentials and prediction errors. We investigated the neural basis of such hierarchical expectations of semantics in a word-pair priming paradigm. By manipulating the overall proportion of related or unrelated word-pairs across the task, we created two global contexts that differentially encouraged strategic use of primes. Across two experiments, we replicated behavioural evidence of greater priming in the high validity context, reflecting strategic expectations of upcoming targets based on ‘global’ context. In our pre-registered EEG analyses, we observed a ‘local’ prediction error ERP effect (i.e. semantic priming) approximately 250ms post-target, which, in exploratory analyses, was followed 100ms later by a signal that interacted with the global context. However, the later effect behaved in an apredictive manner - i.e. was most extreme for fulfilled expectations, rather than violations. Our results are consistent with interpretations of early ERPs as reflections of prediction error and later ERPs as processes related to conscious access and in support of task demands. Significance statement Semantic expectations have been associated with the ERP N400 component, which is modulated by semantic prediction errors across levels of the hierarchy. However, there is no evidence of a two-stage profile that reflects violations of semantic expectations at a single level of the hierarchy, such as the MMN and P3b observed in the local-global paradigm, which are elicited by violations of local and global expectations, respectively. In the present study, we provided evidence of an early ERP effect that reflects violations of local semantic expectations, followed by an apredictive signal that interacted with the global context. Thus, these results support the notion of early ERPs as prediction errors and later ERPs reflecting conscious access and strategic use of context.
Comprehension of degraded speech requires higher-order expectations informed by prior knowledge. Accurate top-down expectations of incoming degraded speech cause a subjective semantic 'pop-out' or conscious breakthrough experience. Indeed, the same stimulus can be perceived as meaningless when no expectations are made in advance. We investigated the event-related potential (ERP) correlates of these top-down expectations, their error signals and the subjective pop-out experience in healthy participants. We manipulated expectations in a word-pair priming degraded (noise-vocoded) speech task and investigated the role of top-down expectation with a between-groups attention manipulation. Consistent with the role of expectations in comprehension, repetition priming significantly enhanced perceptual intelligibility of the noise-vocoded degraded targets for attentive participants. An early ERP was larger for mismatched (i.e. unexpected) targets than matched targets, indicative of an initial error signal not reliant on top-down expectations. Subsequently, a P3a-like ERP was larger to matched targets than mismatched targets only for attending participants-i.e. a pop-out effect-while a later ERP was larger for mismatched targets and did not significantly interact with attention. Rather than relying on complex post hoc interactions between prediction error and precision to explain this apredictive pattern, we consider our data to be consistent with prediction error minimization accounts for early stages of processing followed by Global Neuronal Workspace-like breakthrough and processing in service of task goals.
Functional MRI at ultra-high field (UHF, ≥7T) provides significant increases in BOLD contrast-to-noise ratio (CNR) compared with conventional field strength (3T), and has been exploited for reduced field-of-view, high spatial resolution mapping of primary sensory areas. Applying these high spatial resolution methods to investigate whole brain functional responses to higher-order cognitive tasks leads to a number of challenges, in particular how to perform robust group-level statistical analyses. This study addresses these challenges using an inter-sensory cognitive task which modulates top-down attention at graded levels between the visual and somatosensory domains. At the individual level, highly focal functional activation to the task and task difficulty (modulated by attention levels) were detectable due to the high CNR at UHF. However, to assess group level effects, both anatomical and functional variability must be considered during analysis. We demonstrate the importance of surface over volume normalization and the requirement of no spatial smoothing when assessing highly focal activity. Using novel group analysis on anatomically parcellated brain regions, we show that in higher cognitive areas (parietal and dorsal-lateral-prefrontal cortex) fMRI responses to graded attention levels were modulated quadratically, whilst in visual cortex and VIP, responses were modulated linearly. These group fMRI responses were not seen clearly using conventional second-level GLM analyses, illustrating the limitations of a conventional approach when investigating such focal responses in higher cognitive regions which are more anatomically variable. The approaches demonstrated here complement other advanced analysis methods such as multi-variate pattern analysis, allowing UHF to be fully exploited in cognitive neuroscience.
Directing attention helps to extract relevant information and suppress distracters. Alpha brain oscillations (8-12 Hz) are crucial for this process, with power decreases facilitating processing of important information and power increases inhibiting brain regions processing irrelevant information. Evidence for this phenomenon arises from visual attention studies (Worden et al., 2000); however, the effect also exists in other modalities, including the somatosensory system (Haegens et al., 2011) and intersensory attention tasks (Foxe and Snyder, 2011). We investigated in human participants (10 females, 10 males) the role of alpha oscillations in focused (0/100%) versus divided (40/60%) attention, both across modalities (visual/somatosensory; Experiment 1) and within the same modality (visual domain: across hemifields; Experiment 2) while recording EEG over 128 scalp electrodes. In Experiment 1, participants divided their attention between visual and somatosensory modality to determine the temporal/spatial frequency of a target stimulus (vibrotactile stimulus/Gabor grating). In Experiment 2, participants divided attention between two visual hemifields to identify the orientation of a Gabor grating. In both experiments, prestimulus alpha power in visual areas decreased linearly with increasing attention to visual stimuli. In contrast, prestimulus alpha power in parietal areas was lower when attention was divided between modalities/hemifields compared with focused attention. These results suggest there are two alpha sources, one of which reflects the "visual spotlight of attention" and the other reflects attentional effort. To our knowledge, this is the first study to show that attention recruits two spatially distinct alpha sources in occipital and parietal brain regions, acting simultaneously but serving different functions in attention.SIGNIFICANCE STATEMENT Attention to one spatial location/sensory modality leads to power changes of alpha oscillations (∼10 Hz) with decreased power over regions processing relevant information and power increases to actively inhibit areas processing "to-be-ignored" information. Here, we used detailed source modeling to investigate EEG data recorded during separate unimodal (visual) and multimodal (visual and somatosensory) attention tasks. Participants either focused their attention on one modality/spatial location or directed it to both. We show for the first time two distinct alpha sources are active simultaneously but play different roles. A sensory (visual) alpha source was linearly modulated by attention representing the "visual spotlight of attention." By contrast, a parietal alpha source was modulated by attentional effort, showing lowest alpha power when attention was divided.
The notion of semantic embodiment posits that concepts are represented in the same neural sensorimotor systems that were involved in their acquisition. However, evidence in support of embodied semantics – in particular the hypothesised contribution of motor and premotor cortex to the representation of action concepts – is varied. Here, we tested the hypothesis that, consistent with semantic embodiment, sensorimotor cortices will rapidly become active while healthy participants access the meaning of visually-presented motor and non-motor action verbs. Event-related potentials revealed early differential processing of motor and non-motor verbs (164–203 ms) within distinct regions of cortex likely reflecting rapid cortical activation of differentially distributed semantic representations. However, we found no evidence for a specific role of sensorimotor cortices in supporting these representations. Moreover, we observed a later modulation of the alpha band (8–12 Hz) from 555–785 ms over central electrodes, with estimated generators within the left superior parietal lobule, which may reflect post-lexical activation of the object-directed features of the motor action concepts. In conclusion, we find no evidence for a specific role of sensorimotor cortices when healthy participants judge the meaning of visually-presented action verbs. However, the relative contribution of sensorimotor cortices to action comprehension may vary as a function of task goals.
Background Transcranial alternating current stimulation (tACS) is widely used to entrain or modulate brain oscillations in order to investigate causal relationships between oscillations and cognition. Objective In a series of experiments we here addressed the question of whether event-related, transient tACS in the beta frequency range can be used to entrain beta oscillations in two different domains: episodic memory formation and motor cortex excitability. Methods In experiments 1 and 2, 72 healthy human participants engaged in an incidental encoding task of verbal and non-verbal material while receiving tACS to the left and right inferior frontal gyrus (IFG) at 6.8Hz, 10.7Hz, 18.5Hz, 30Hz, 48Hz and sham stimulation for 2s during stimulus presentation. In experiment 3, tACS was administered to M1 at the individual motor beta frequency of eight subjects. We investigated the relationship between the size of TMS induced MEPs and tACS phase. Results Beta tACS did not affect memory performance compared to sham stimulation in experiments 1 and 2. Likewise, in experiment 3, MEP size was not modulated by the tACS phase. Conclusions Our findings suggest that event-related, transient tACS in the beta frequency range cannot be used to modulate the formation of episodic memories or motor cortex excitability. These null-results question the effectiveness of event-related tACS to entrain beta oscillations and modulate cognition.
Ongoing brain oscillations (7–10 Hz) modulate visual perception; in particular, their precise phase can predict target perception. Here, we employ this phase-dependence of perception in a psychophysical experiment to track spatial properties of entrained oscillations of visual perception across the visual field. Is this entrainment local, or a more global phenomenon? If the latter, does oscillatory phase synchronize over space, or vary with increasing distance from the oscillatory source? We presented a disc stimulus in the upper left quadrant, oscillating in luminance at different frequencies (individual alpha frequency (IAF), 5 Hz and 15 Hz) to entrain an oscillation with specific frequency and spatial origin. Observers fixated centrally, while flash stimuli at perceptual threshold appeared at different positions and times with respect to the oscillating stimulus. IAF and 5 Hz luminance oscillations modulated detection performance at all tested positions, whereas at 15 Hz, the effect was weaker and less consistent. Furthermore, for IAF and 5 Hz entrainment, preferred phases for target detection differed significantly between spatial locations, suggesting “local” entrainment of detection performance next to the oscillatory source, whereas more distant target locations shared a “global” effect with a significantly different phase. This unexpected global component of entrainment is tentatively attributed to widespread connectivity from thalamic nuclei such as the pulvinar.
Il a ete propose que la perception visuelle soit organisee de facon discrete, reposant sur l'activite cerebrale oscillatoire d'environ 10Hz. Bien qu'en condition normale notre perception visuelle nous paraisse continue, nous montrons dans une premiere experience que cette continuite peut etre interrompue lors de la perception d'un stimulus specifique pourtant statique, revelant les cycles d'oscillations alpha (~10Hz) a la base de la perception visuelle. La 'Flickering Wheel Illusion' ('L'illusion de la roue qui clignote') genere un clignotement illusoire regulier, reapparaissant toutes les 100ms ce qui correspond a une phase du cycle alpha. Des etudes precedentes ont montre que certaines phases des oscillations alpha occipitales (tout comme des oscillations theta fronto-centrales) portent des proprietes 'inhibitrices' en vue du traitement visuel tandis que leurs phases opposees tendent a augmenter la probabilite de detecter des stimuli visuels. Dans une deuxieme etude, nous avons analyse comment ce rapport phase-detection temporel est represente au niveau spatial dans le cortex visuel. Pour cela, nous avons entraine des oscillations alpha via le clignotement d'un disque a une certaine position spatiale du champ visuel, pendant que les participants avaient pour tâche de detecter des stimuli visuels de basse luminance pouvant apparaitre a de multiples positions spatiales. Les oscillations entrainees modulaient la performance de detection a toutes les positions auxquelles des cibles pourraient apparaitre montrant des maxima a differentes phases de l'oscillation. De facon interessante, nous avons pu montrer que la phase des oscillations entrainees se propageait dans l'espace comme une onde progressive, une propriete qui pourrait impliquer des roles fonctionnels specifiques pour le traitement sensoriel. En plus de son influence sur les processus cognitives comme la perception visuelle, la phase des oscillations lentes peut aussi moduler l'amplitude d'une oscillation rapide ('phase-amplitude cross-frequency coupling' ; 'couplage phase-amplitude inter-frequentiel' ;'PAC'). Dans une troisieme experience, nous avons revele l'existence d'un PAC entre des oscillations alpha et gamma pendant l'etat de repos (en l'absence de tâche cognitive specifique) ce que nous avons assigne a de 'l'inhibition pulsative', un mecanisme de controle du traitement sensoriel porte par le rythme alpha. Selon cette theorie, les oscillations alpha mettraient donc a disposition des moments ou bien des phases specifiques qui menent a une augmentation d'amplitude d'oscillations gamma entrainant une augmentation du traitement sensoriel, tandis que des phases opposees inhiberaient le traitement sensoriel. En outre, nous avons observe un PAC robuste entre des oscillations theta et beta pendant l'etat de repos ainsi que pendant une tâche de detection. Ce PAC pourrait aider a maintenir un etat d'activite cerebrale de base. Finalement, nous avons pu reveler un PAC entre des oscillations theta et gamma principalement dans des regions fronto-centrales, influencant la perception visuelle avec emploi d'attention visuelle. Ce resultat est en continuite et complete des resultats precedents de notre groupe qui montraient l'influence de la phase des oscillations theta sur la perception visuelle. Le travail presente dans cette these contribue aux connaissances du role important des oscillations spontanees pour la perception visuelle en utilisant plusieurs approches experimentales et revelent de nouvelles questions de ce champ de recherche.