Michael A. Pitts

Isolating neural correlates of conscious perception from neural correlates of reporting one's perception

To isolate neural correlates of conscious perception (NCCs), a standard approach has been to contrast neural activity elicited by identical stimuli of which subjects are aware vs. unaware. Because conscious experience is private, determining whether a stimulus was consciously perceived requires subjective report: e.g., button-presses indicating detection, visibility ratings, verbal reports, etc. This reporting requirement introduces a methodological confound when attempting to isolate NCCs: The neural processes responsible for accessing and reporting ones percept are difficult to distinguish from those underlying the conscious percept itself. Here, we review recent attempts to circumvent this issue via a modified inattentional blindness paradigm (Pitts et al., 2012) and present new data from a backward masking experiment in which task-relevance and visual awareness were manipulated in a 2 × 2 crossed design. In agreement with our previous inattentional blindness results, stimuli that were consciously perceived yet not immediately accessed for report (aware, task-irrelevant condition) elicited a mid-latency posterior ERP negativity (~200–240 ms), while stimuli that were accessed for report (aware, task-relevant condition) elicited additional components including a robust P3b (~380–480 ms) subsequent to the mid-latency negativity. Overall, these results suggest that some of the NCCs identified in previous studies may be more closely linked with accessing and maintaining perceptual information for reporting purposes than with encoding the conscious percept itself. An open question is whether the remaining NCC candidate (the ERP negativity at 200–240 ms) reflects visual awareness or object-based attention.

Right Parietal Brain Activity Precedes Perceptual Alternation During Binocular Rivalry

We investigated perceptual reversals for intermittently presented stimuli during binocular rivalry and physical alternation while the ongoing EEG was recorded from 64 channels. EEG topographies immediately preceding stimulus‐onset were analyzed and two topographies doubly dissociated perceptual reversals from non‐reversals. The estimated intracranial generators associated with these topographies were stronger in right inferior parietal cortex and weaker bilaterally in the ventral stream before perceptual reversals. No such differences were found for physical alternation of the same stimuli. These results replicate and extend findings from a previous study with the Necker cube and suggest common neural mechanisms associated with perceptual reversals during binocular rivalry and ambiguous figure perception. For both types of multi‐stable stimuli, the dorsal stream is more active preceding perceptual reversals. Activity in the ventral stream, however, differed for binocular rivalry compared to ambiguous figures. The results from the two studies suggest a causal role for the right inferior parietal cortex in generating perceptual reversals regardless of the type of multi‐stable stimulus, while activity in the ventral stream appears to depend on the particular type of stimulus. Hum Brain Mapp 32:1432–1442, 2010.

Insights from Intermittent Binocular Rivalry and EEG

Novel stimulation and analytical approaches employed in EEG studies of ambiguous figures have recently been applied to binocular rivalry. The combination of intermittent stimulus presentation and EEG source imaging has begun to shed new light on the neural underpinnings of binocular rivalry. Here, we review the basics of the intermittent paradigm and highlight methodological issues important for interpreting previous results and designing future experiments. We then outline current analytical approaches, including EEG microstates, event-related potentials, and statistically based source estimation, and propose a neural model of the sequence of brain events that may underlie different aspects of binocular rivalry. Finally, we discuss the advantages and limitations of using binocular rivalry as a tool to investigate the neural basis of perceptual awareness.

Neural generators of ERPs linked with Necker cube reversals

Multistable perception occurs when a single physical stimulus leads to two or more distinct percepts that spontaneously switch (reverse). Previous ERP studies have reported reversal negativities and late positive components associated with perceptual reversals. The goal of the current study was to localize the neural generators of the reversal ERP components in order to evaluate their correspondence with previous fMRI results and to better understand their functional significance. A Necker-type stimulus was presented for brief intervals while subjects indicated their perceptions. Local auto-regressive average source analyses and dipole modeling indicated that sources for the reversal negativity were located in inferior occipital-temporal cortex. Generators of the late positive component were estimated to reside in inferior temporal and superior parietal regions.

Perceptual reversals during binocular rivalry: ERP components and their concomitant source differences

We used an intermittent stimulus presentation to investigate event-related potential (ERP) components associated with perceptual reversals during binocular rivalry. The combination of spatiotemporal ERP analysis with source imaging and statistical parametric mapping of the concomitant source differences yielded differences in three time windows: reversals showed increased activity in early visual (∼120 ms) and in inferior frontal and anterior temporal areas (∼400-600 ms) and decreased activity in the ventral stream (∼250-350 ms). The combination of source imaging and statistical parametric mapping suggests that these differences were due to differences in generator strength and not generator configuration, unlike the initiation of reversals in right inferior parietal areas. These results are discussed within the context of the extensive network of brain areas that has been implicated in the initiation, implementation, and appraisal of bistable perceptual reversals.

Neural Signatures of Conscious Face Perception in an Inattentional Blindness Paradigm

Previous studies suggest that early stages of face-specific processing are performed preattentively and unconsciously, whereas conscious perception emerges with late-stage (>300 ms) neuronal activity. A conflicting view, however, posits that attention is necessary for face-specific processing and that early-to-mid latency neural responses (∼100–300 ms) correspond more closely with perceptual awareness. The current study capitalized on a recently developed method for manipulating attention and conscious perception during EEG recording (modified inattentional blindness paradigm) and used face stimuli that elicit a well known marker of early face processing, the N170 event-related potential (ERP). In Phase 1 of the experiment, subjects performed a demanding distracter task while line drawings of faces and matched control stimuli were presented in the center of their view. When queried, half of the subjects reported no awareness of the faces and were deemed inattentionally blind. In Phase 2, subjects performed the same distracter task, but now consciously perceived the face stimuli due to the intervening questioning. In Phase 3, subjects performed a discrimination task on the faces. Two primary contrasts were made: aware versus unaware (equally task irrelevant) and task-relevant versus task-irrelevant (equally aware). The N170 and a subsequent ERP component, the visual awareness negativity (∼260–300 ms), were absent during inattentional blindness and present in the aware conditions. The P3b (>300 ms) was absent for task-irrelevant faces, even when consciously perceived, and present only when the faces were task relevant. These results inform contemporary theories of conscious face perception in particular and visual attention and perceptual awareness in general.

Perception of ensemble statistics requires attention

To overcome inherent limitations in perceptual bandwidth, many aspects of the visual world are represented as summary statistics (e.g., average size, orientation, or density of objects). Here, we investigated the relationship between summary (ensemble) statistics and visual attention. Recently, it was claimed that one ensemble statistic in particular, color diversity, can be perceived without focal attention. However, a broader debate exists over the attentional requirements of conscious perception, and it is possible that some form of attention is necessary for ensemble perception. To test this idea, we employed a modified inattentional blindness paradigm and found that multiple types of summary statistics (color and size) often go unnoticed without attention. In addition, we found attentional costs in dual-task situations, further implicating a role for attention in statistical perception. Overall, we conclude that while visual ensembles may be processed efficiently, some amount of attention is necessary for conscious perception of ensemble statistics.

Testing domain-general theories of perceptual awareness with auditory brain responses

Past research has identified several candidate neural correlates of consciousness (NCCs) during visual perception. Recent research on auditory perception shows promise for establishing the generality of various NCCs across sensory modalities, as well as for revealing differences in how conscious processing unfolds in different sensory systems.

Chromatic perceptive field sizes change with retinal illuminance

The effect of retinal illuminance (0.3-3.3 log td) on chromatic perceptive field size was investigated at 10 degrees eccentricity along the horizontal meridian of the temporal retina. Using the 4+1 color-naming procedure, observers described the hue and saturation of a series of monochromatic stimuli (440-660 nm, in 10-nm steps) of various test sizes (.098-5 degrees) after 30-min dark adaptation. Perceptive field sizes of the four elemental hues and the saturation component were estimated for each wavelength at each retinal illuminance. Results indicate that perceptive field sizes for blue, green, yellow, and saturation all decrease with increasing retinal illuminance; the perceptive field size for red is the smallest and invariant with intensity. The influence of rods on perceptive field size may account for some of the results; other factors are also considered.

Effect of stimulus intensity on the sizes of chromatic perceptive fields.

The effects of intensity on chromatic perceptive field size were investigated along the horizontal meridian at 10 degrees temporal eccentricity by manipulating stimulus intensity from 0.3 to 3.3 log trolands. Following light adaptation, observers described the hue and saturation of monochromatic stimuli (440-660 nm, in 10 nm steps) for a series of test sizes (0.098-3 degrees) presented along the time period associated with the cone plateau of the dark-adaptation function. Perceptive field sizes of the four elemental hues (red, green, yellow, and blue) and the saturation component were estimated by three observers at each intensity level for each wavelength. In general, perceptive field sizes of blue and red are the smallest, and yellow and green are the largest. Furthermore, perceptive field sizes of all four hues decrease with increasing stimulus intensity, though the absolute change is largest for green and yellow. The decrease in size with increase in intensity cannot be completely explained in terms of saturation or rod signals and is likely, then, attributable to a cone-based mechanism.