We perceive visual objects as unified although different brain areas process different features. An attentional mechanism has been proposed to be involved with feature binding, as evidenced by observations of binding errors (i.e., illusory conjunctions) when attention is diverted. However, the neural underpinnings of this feature binding are not well understood. We examined the neural mechanisms of feature binding by recording EEG during an attentionally demanding discrimination task. Unlike prestimulus alpha oscillatory activity and early ERPs (i.e., the N1 and P1 components), the N1pc, reflecting stimulus-evoked spatial attention, was reduced for errors relative to correct responses and illusory conjunctions. However, the later SPCN, reflecting visual short-term memory, was reduced for illusory conjunctions and errors compared with correct responses. Furthermore, binding errors were associated with distinct posterior lateralized activity during this 200- to 300-msec window. These results implicate a temporal binding window that integrates visual features after stimulus-evoked attention but before encoding into visual short-term memory.

Variability in perception between individuals may be a consequence of different inherent neural processing speeds. To assess whether alpha oscillations systematically reflect a feedback pacing mechanism for cortical processing during visual perception, comparisons were made between alpha oscillations, visual suppression from TMS, visual evoked responses, and metacontrast masking. Peak alpha oscillation frequencies, measured through scalp EEG recordings, significantly correlated with the optimum latencies for visual suppression from TMS of early visual cortex. Individuals with shorter alpha periods (i.e., higher peak alpha frequencies) processed visual information faster than those with longer alpha periods (i.e., lower peak alpha frequencies). Moreover, peak alpha oscillation periods and optimum TMS visual suppression latencies predicted the latencies of late but not early visual evoked responses. Together, these findings demonstrate an important role of alpha oscillatory and late feedback activity in visual cortex for conscious perception. They also show that the timing for visual awareness varies across individuals, depending on the pace of one's endogenous oscillatory cycling frequency.

We investigated the dynamics of brain processes facilitating conscious experience of external stimuli. Previously, we proposed that alpha (8–12 Hz) oscillations, which fluctuate with both sustained and directed attention, represent a pulsed inhibition of ongoing sensory brain activity. Here we tested the prediction that inhibitory alpha oscillations in visual cortex are modulated by top–down signals from frontoparietal attention networks. We measured modulations in phase-coherent alpha oscillations from superficial frontal, parietal, and occipital cortices using the event-related optical signal (EROS), a measure of neuronal activity affording high spatiotemporal resolution, along with concurrently recorded EEG, while participants performed a visual target detection task. The pretarget alpha oscillations measured with EEG and EROS from posterior areas were larger for subsequently undetected targets, supporting alpha's inhibitory role. Using EROS, we localized brain correlates of these awareness-related alpha oscillations measured at the scalp to the cuneus and precuneus. Crucially, EROS alpha suppression correlated with posterior EEG alpha power across participants. Sorting the EROS data based on EEG alpha power quartiles to investigate alpha modulators revealed that suppression of posterior alpha was preceded by increased activity in regions of the dorsal attention network and decreased activity in regions of the cingulo-opercular network. Cross-correlations revealed the temporal dynamics of activity within these preparatory networks before posterior alpha modulation. The novel combination of EEG and EROS afforded localization of the sources and correlates of alpha oscillations and their temporal relationships, supporting our proposal that top–down control from attention networks modulates both posterior alpha and awareness of visual stimuli.

We examined the causal relationship between the phase of alpha oscillations (9–12 Hz) and conscious visual perception using rhythmic TMS (rTMS) while simultaneously recording EEG activity. rTMS of posterior parietal cortex at an alpha frequency (10 Hz), but not occipital or sham rTMS, both entrained the phase of subsequent alpha oscillatory activity and produced a phase-dependent change on subsequent visual perception, with lower discrimination accuracy for targets presented at one phase of the alpha oscillatory waveform than for targets presented at the opposite phase. By extrinsically manipulating the phase of alpha before stimulus presentation, we provide direct evidence that the neural circuitry in the parietal cortex involved with generating alpha oscillations plays a causal role in determining whether or not a visual stimulus is successfully perceived.

Although examples of unconscious shape priming have been well documented, whether such priming requires early visual cortex (V1/V2) has not been established. In the current study, we used TMS of V1/V2 at varying temporal intervals to suppress the visibility of preceding shape primes while the interval between primes and targets was kept constant. Our results show that, although conscious perception requires V1/V2, unconscious priming can occur without V1/V2 at an intermediate temporal interval but not at early (5–25 msec) or later (65–125 msec) stages of processing. Because the later time window of unconscious priming suppression has been proposed to interfere with feedback processing, our results further suggest that feedback processing is also essential for unconscious priming and may not be a sufficient condition for conscious vision.

Unconscious processing has been convincingly demonstrated for task-relevant feature dimensions. However, it is possible that the visual system is capable of more complex unconscious operations, extracting visual features even when they are unattended and task irrelevant. In the current study, we addressed this question by measuring unconscious priming using a task in which human participants attended to a target object's shape while ignoring its color. We measured both behavioral priming effects and priming-related fMRI activations from primes that were unconsciously presented using metacontrast masking. The results showed faster RTs and decreases in fMRI activation only when the primes were identical to the targets, indicating that primes were processed both in the attended shape and the unattended color dimensions. Reductions in activation were observed in early visual areas, including primary visual cortex, as well as in feature-responsive areas for shape and color. These results indicate that multiple features can be unconsciously encoded and possibly bound using the same visual networks activated by consciously perceived images.

Visual stability refers to our stable visuospatial perceptions despite the unstable visual input caused by saccades. Functional neuroimaging results, studies on patients with posterior parietal cortex (PPC) lesions, and single-unit recordings in the lateral intraparietal sulcus of primates indirectly suggest that the PPC might be a potential locus of visual stability through its involvement with spatial remapping. Here we directly explored the role of the PPC in visual stability by applying transcranial magnetic stimulation (TMS) while participants performed a perisaccadic displacement detection task. We show that TMS over the PPC but not a frontal control site alters sensitivity to displacement detection when administered just before contralateral saccades and that a general impairment in attention or in the perception of apparent motion cannot account for the decreased sensitivity. The specific relationship between the timing of TMS and saccade direction demonstrates that saccadic suppression of displacement (SSD) is likely a consequence of noisy contralateral spatial representations in the PPC around the time of a saccade. The same mechanism may keep the unstable visual world in the temporal proximity of saccades from reaching our consciousness.

Rice University The visual modality typically dominates over our other senses. Here we show that after inducing an extreme conflict in the left hand between vision of touch (present) and the feeling of touch (absent), sensitivity to touch increases for several minutes after the conflict. Transcranial magnetic stimulation of the posterior parietal cortex after this conflict not only eliminated the enduring visual enhancement of touch, but also impaired normal tactile perception. This latter finding demonstrates a direct role of the parietal lobe in modulating tactile perception as a result of the conflict between these senses. These results provide evidence for visual-to-tactile perceptual modulation and demonstrate effects of illusory vision of touch on touch perception through a long-lasting modulatory process in the posterior parietal cortex.

We examined the effects of chronic unilateral lesions to either the inferior parietal lobe, or to the dorsolateral prefrontal cortex including the frontal eye fields (FEFs), upon human visual perception and saccades in temporal-order-judgment (TOJ) tasks. Two visual events were presented on each trial, one in each hemifield at various stimulus onset asynchronies (SOAs). In the saccade task, patients moved their eyes to whichever stimulus attracted gaze first. In the perceptual-manual task, they pressed a button to indicate which stimulus was perceived first. Frontal patients showed appropriate TOJs for visual targets in both tasks. Parietal patients showed appropriate TOJs in the perceptual-manual but not the saccade task; their saccades tended to be ipsilesional unless the contralesional target led substantially. This reveals a bias in saccade choice after parietal damage that cannot be attributed to deficient visual perception. These results challenge previous claims that only anterior lesions produce motoric spatial biases in humans. However, they are in accord with recent neurophysiological evidence for parietal involvement in saccade generation, and also with suggestions that visuomotor transformations in the parietal lobe serving direct spatial motor responses can dissociate from conscious perception as indicated by indirect arbitrary responses.

The contributions of the superior prefrontal cortex (SPFC) and the superior parietal lobule (SPL) in generating voluntary endogenous and reflexive visually guided saccades were investigated using transcranial magnetic stimulation (TMS). Subjects made choice saccades to the left or right visual field in response to a central arrowhead (endogenous go signal) or a peripheral asterisk (exogenous go signal) that were presented along with a single TMS pulse at varying temporal intervals. TMS over the SPFC increased latencies for saccades made in response to an endogenous go signal toward the contralateral hemifield. No effects were observed when the go signal was exogenous and TMS was over the SPFC or when TMS was over the SPL for either saccade type. The delayed contralateral endogenous saccades observed in this study are likely a consequence of disruption in the normal operations of the human frontal eye field.