Tentative fMRI signatures of perceptual echoes in early visual cortex.

The visual Impulse Response Function (IRF) can be estimated by cross-correlating random luminance sequences with concurrently recorded EEG. It typically contains a strong 10 Hz oscillatory component, suggesting that visual information reverberates in the human brain as a "perceptual echo". The neural origin of these echoes remains unknown. To address this question, we recorded EEG and fMRI in two separate sessions. In both sessions, a disk whose luminance followed a random (white noise) sequence was presented in the upper left quadrant. Individual IRFs were derived from the EEG session. These IRFs were then used as "response templates" to reconstruct an estimate of the EEG during the fMRI session, by convolution with the corresponding random luminance sequences. The 7-14 Hz (alpha, the main frequency component of the IRF) envelope of the reconstructed EEG was finally used as an fMRI regressor, to determine which brain voxels co-varied with the oscillations elicited by the luminance sequence, i.e., the "perceptual echoes". The reconstructed envelope of EEG alpha was significantly correlated with BOLD responses in V1 and V2. Surprisingly, this correlation was visible outside, but not within the directly (retinotopically) stimulated region. We tentatively interpret this lack of alpha modulation as a BOLD saturation effect, since the overall stimulus-induced BOLD response was inversely related, across voxels, to the signal variability over time. In conclusion, our results suggest that perceptual echoes originate in early visual cortex, driven by widespread activity in V1 and V2, not retinotopically restricted, but possibly reflecting the propagation of a travelling alpha wave.

Alpha Power Modulates Perception Independently of Endogenous Factors.

Oscillations are ubiquitous in the brain. Alpha oscillations in particular have been proposed to play an important role in sensory perception. Past studies have shown that the power of ongoing EEG oscillations in the alpha band is negatively correlated with visual outcome. Moreover, it also co-varies with other endogenous factors such as attention, vigilance, or alertness. In turn, these endogenous factors influence visual perception. Therefore, it remains unclear how much of the relation between alpha and perception is indirectly mediated by such endogenous factors, and how much reflects a direct causal influence of alpha rhythms on sensory neural processing. We propose to disentangle the direct from the indirect causal routes by introducing modulations of alpha power, independently of any fluctuations in endogenous factors. To this end, we use white-noise sequences to constrain the brain activity of 20 participants. The cross-correlation between the white-noise sequences and the concurrently recorded EEG reveals the impulse response function (IRF), a model of the systematic relationship between stimulation and brain response. These IRFs are then used to reconstruct rather than record the brain activity linked with new random sequences (by convolution). Interestingly, this reconstructed EEG only contains information about oscillations directly linked to the white-noise stimulation; fluctuations in attention and other endogenous factors may still modulate brain alpha rhythms during the task, but our reconstructed EEG is immune to these factors. We found that the detection of near-perceptual threshold targets embedded within these new white-noise sequences depended on the power of the ~10 Hz reconstructed EEG over parieto-occipital channels. Around the time of presentation, higher power led to poorer performance. Thus, fluctuations in alpha power, induced here by random luminance sequences, can directly influence perception: the relation between alpha power and perception is not a mere consequence of fluctuations in endogenous factors.

At What Latency Does the Phase of Brain Oscillations Influence Perception?

Recent evidence has shown a rhythmic modulation of perception: prestimulus ongoing electroencephalography (EEG) phase in the θ (4-8 Hz) and α (8-13 Hz) bands has been directly linked with fluctuations in target detection. In fact, the ongoing EEG phase directly reflects cortical excitability: it acts as a gating mechanism for information flow at the neuronal level. Consequently, the key phase modulating perception should be the one present in the brain when the stimulus is actually being processed. Most previous studies, however, reported phase modulation peaking 100 ms or more before target onset. To explain this discrepancy, we first use simulations showing that contamination of spontaneous oscillatory signals by target-evoked ERP and signal filtering (e.g., wavelet) can result in an apparent shift of the peak phase modulation towards earlier latencies, potentially reaching the prestimulus period. We then present a paradigm based on linear systems analysis which can uncover the true latency at which ongoing EEG phase influences perception. After measuring the impulse response function, we use it to reconstruct (rather than record) the brain activity of human observers during white noise sequences. We can then present targets in those sequences, and reliably estimate EEG phase around these targets without any influence of the target-evoked response. We find that in these reconstructed signals, the important phase for perception is that of fronto-occipital ∼6 Hz background oscillations at about 75 ms after target onset. These results confirm the causal influence of phase on perception at the time the stimulus is effectively processed in the brain.

The ability of the auditory system to cope with temporal subsampling depends on the hierarchical level of processing.

Evidence for rhythmic or 'discrete' sensory processing is abundant for the visual system, but sparse and inconsistent for the auditory system. Fundamental differences in the nature of visual and auditory inputs might account for this discrepancy: whereas the visual system mainly relies on spatial information, time might be the most important factor for the auditory system. In contrast to vision, temporal subsampling (i.e. taking 'snapshots') of the auditory input stream might thus prove detrimental for the brain as essential information would be lost. Rather than embracing the view of a continuous auditory processing, we recently proposed that discrete 'perceptual cycles' might exist in the auditory system, but on a hierarchically higher level of processing, involving temporally more stable features. This proposal leads to the prediction that the auditory system would be more robust to temporal subsampling when applied on a 'high-level' decomposition of auditory signals. To test this prediction, we constructed speech stimuli that were subsampled at different frequencies, either at the input level (following a wavelet transform) or at the level of auditory features (on the basis of LPC vocoding), and presented them to human listeners. Auditory recognition was significantly more robust to subsampling in the latter case, that is on a relatively high level of auditory processing. Although our results do not directly demonstrate perceptual cycles in the auditory domain, they (a) show that their existence is possible without disrupting temporal information to a critical extent and (b) confirm our proposal that, if they do exist, they should operate on a higher level of auditory processing.