Barbara F. Händel

Alpha oscillations correlate with the successful inhibition of unattended stimuli.

Because the human visual system is continually being bombarded with inputs, it is necessary to have effective mechanisms for filtering out irrelevant information. This is partly achieved by the allocation of attention, allowing the visual system to process relevant input while blocking out irrelevant input. What is the physiological substrate of attentional allocation? It has been proposed that alpha activity reflects functional inhibition. Here we asked if inhibition by alpha oscillations has behavioral consequences for suppressing the perception of unattended input. To this end, we investigated the influence of alpha activity on motion processing in two attentional conditions using magneto-encephalography. The visual stimuli used consisted of two random-dot kinematograms presented simultaneously to the left and right visual hemifields. Subjects were cued to covertly attend the left or right kinematogram. After 1.5 sec, a second cue tested whether subjects could report the direction of coherent motion in the attended (80%) or unattended hemifield (20%). Occipital alpha power was higher contralateral to the unattended side than to the attended side, thus suggesting inhibition of the unattended hemifield. Our key finding is that this alpha lateralization in the 20% invalidly cued trials did correlate with the perception of motion direction: Subjects with pronounced alpha lateralization were worse at detecting motion direction in the unattended hemifield. In contrast, lateralization did not correlate with visual discrimination in the attended visual hemifield. Our findings emphasize the suppressive nature of alpha oscillations and suggest that processing of inputs outside the field of attention is weakened by means of increased alpha activity.

Cross-frequency coupling of brain oscillations indicates the success in visual motion discrimination

Cortical activity such as recorded by EEG or MEG is characterized by ongoing rhythms that encompass a wide range of temporal and spatial scales. Recent studies have suggested an oscillatory hierarchy with faster oscillations being locked to preferred phases of underlying slower waves, a functional principle applied up to the level of action potential generation. We here tested the idea that amplitude-phase coupling between frequencies might serve the detection of weak sensory signals. To this end we recorded neuromagnetic responses during a motion discrimination task using near-threshold stimuli. Amplitude modulation of occipital high-frequency oscillations in the gamma range (63+/-5 Hz) was phase locked to a slow-frequency oscillation in the delta band (1-5 Hz). Most importantly, the strength of gamma amplitude modulation reflected the success in visual discrimination. This correlation provides evidence for the hypothesis that coupling between low- and high-frequency brain oscillations subserves signal detection.

Visual Motion Perception Deficits Due to Cerebellar Lesions Are Paralleled by Specific Changes in Cerebro-Cortical Activity

Recent anatomical studies have revealed strong cerebellar projections into parietal and prefrontal cortex. These findings suggest that the cerebellum might not only play a functional role in motor control but also cognitive domains, an idea also supported by neuropsychological testing of patients with cerebellar lesions that has revealed specific deficits. The goal of the present study was to test whether or not cognitive impairments after cerebellar damage are resulting from changes in cerebro-cortical signal processing. The detection of global visual motion embedded in noise, a faculty compromised after cerebellar lesions, was chosen as a model system. Using magnetoencephalography, cortical responses were recorded in a group of patients with cerebellar lesions (n = 8) and controls (n = 13) who observed visual motion of varied coherence, i.e., motion strength, presented in the peripheral visual field during controlled stationary fixation. Corroborating earlier results, the patients showed a significant impairment in global motion discrimination despite normal fixation behavior. This deficit was paralleled by qualitative differences in responses recorded from parieto-temporal cortex, including a reduced responsiveness to coherent visual motion and a striking loss of bilateral representations of motion coherence. Moreover, the perceptual thresholds correlated with the cortical representation of motion strength on single subject basis. These results demonstrate that visual motion processing in cerebral cortex critically depends on an intact cerebellum and establish a correlation between cortical activity and impaired visual perception resulting from cerebellar damage.

Selective Attention Increases the Dependency of Cortical Responses on Visual Motion Coherence in Man

Attention improves visual discrimination and consequently allows to discern stimuli with low signal-to-noise ratios that otherwise would remain undetected. We used magnetoencephalography (MEG) to test whether neuromagnetic responses recorded from occipito-temporal cortex, reflecting the size of visual motion signals embedded in noise (motion coherence), would mirror the perceptual changes induced by attention. Attention directed to a given hemifield increased and decreased the coherence modulation of the MEG response over contralateral and ipsilateral visual cortex, respectively, indicating a change in the neuronal signal-to-noise ratio at the population level.

Cortical representations of confidence in a visual perceptual decision

To date the exact neuronal implementation of decision confidence has been subject to little research. Here we explore electroencephalographic correlates of human choice certainty in a visual motion discrimination task for either spatial attention or motor effector cue instructions. We demonstrate electrophysiological correlates of choice certainty that evolve as early as 300 ms after stimulus onset and resemble the primary visual motion representations in early visual cortex. These correlates do not emerge unless or until the subject unambiguously knows which of the competing visual stimuli is actually relevant to behaviour. They extend beyond stimulus presentation up to the motor response but are independent of the motor effector. Our findings suggest that perceptual confidence evolves in parallel with representations of stimulus properties and is dedicated to one specific aspect of the visual world. Its electroencephalographic correlates can be disentangled from representations of sensory evidence, objective discrimination performance and overt motor behaviour.

Gamma oscillations underlying the visual motion aftereffect

After having been exposed to strong visual motion in one direction, a subsequently presented stationary visual scene seems to move in the opposite direction. This motion aftereffect (MAE) is usually ascribed to short-term functional changes in cortical areas involved in visual motion analysis akin to adaptation. Using magnetoencephalography (MEG), we show increased global field activity due to the MAE which could mostly be explained by a dipole located near the putative location of human area MT+. We further demonstrate that the induced MAE is accompanied by a significant increase in gamma-band activity (GBA) recorded from parietooccipital cortex contralateral to the visual motion stimulus. This gamma oscillation most likely reflects an increase in neuronal response coherence due to decreased inhibition of a group of neurons with similar preferred direction, namely the direction opposite to the adapted one. A second focal GBA response was picked up by the most posterior sensors ipsilateral to the side of the stimulus, reflecting the size of the MAE, whose source could not be reliably located.

Real color captures attention and overrides spatial cues in grapheme-color synesthetes but not in controls

Grapheme-color synesthetes perceive color when reading letters or digits. We investigated oscillatory brain signals of synesthetes vs. controls using magnetoencephalography. Brain oscillations specifically in the alpha band (∼10Hz) have two interesting features: alpha has been linked to inhibitory processes and can act as a marker for attention. The possible role of reduced inhibition as an underlying cause of synesthesia, as well as the precise role of attention in synesthesia is widely discussed. To assess alpha power effects due to synesthesia, synesthetes as well as matched controls viewed synesthesia-inducing graphemes, colored control graphemes, and non-colored control graphemes while brain activity was recorded. Subjects had to report a color change at the end of each trial which allowed us to assess the strength of synesthesia in each synesthete. Since color (synesthetic or real) might allocate attention we also included an attentional cue in our paradigm which could direct covert attention. In controls the attentional cue always caused a lateralization of alpha power with a contralateral decrease and ipsilateral alpha increase over occipital sensors. In synesthetes, however, the influence of the cue was overruled by color: independent of the attentional cue, alpha power decreased contralateral to the color (synesthetic or real). This indicates that in synesthetes color guides attention. This was confirmed by reaction time effects due to color, i.e. faster RTs for the color side independent of the cue. Finally, the stronger the observed color dependent alpha lateralization, the stronger was the manifestation of synesthesia as measured by congruency effects of synesthetic colors on RTs. Behavioral and imaging results indicate that color induces a location-specific, automatic shift of attention towards color in synesthetes but not in controls. We hypothesize that this mechanism can facilitate coupling of grapheme and color during the development of synesthesia.

Spontaneous local alpha oscillations predict motion-induced blindness

Bistable visual illusions are well suited for exploring the neuronal states of the brain underlying changes in perception. In this study, we investigated oscillatory activity associated with ‘motion‐induced blindness’ (MIB), which denotes the perceptual disappearance of salient target stimuli when a moving pattern is superimposed on them (Bonneh et al., ). We applied an MIB paradigm in which illusory target disappearances would occur independently in the left and right hemifields. Both illusory and real target disappearance were followed by an alpha lateralization with weaker contralateral than ipsilateral alpha activity (~10 Hz). However, only the illusion showed early alpha lateralization in the opposite direction, which preceded the alpha effect present for both conditions and coincided with the estimated onset of the illusion. The duration of the illusory disappearance was further predicted by the magnitude of this early lateralization when considered over subjects. In the gamma band (60–80 Hz), we found an increase in activity contralateral relative to ipsilateral only after a real disappearance. Whereas early alpha activity was predictive of onset and length of the illusory percept, gamma activity showed no modulation in relation to the illusion. Our study demonstrates that the spontaneous changes in visual alpha activity have perceptual consequences.