Endogenous Attention Affects Decision-related Neural Activity But Not Afferent Visual Responses.

Endogenous shifts of spatial attention toward an upcoming stimulus are associated with improvements in behavioral responses to the stimulus, preparatory retinotopic shifts in alpha power, and changes in ERPs. Although attentional modulation of several early sensory ERPs is well established, there is still debate about under what circumstances attention affects the earliest cortical visual evoked response-the C1 ERP component-which is putatively generated from afferent input into primary visual cortex. Moreover, the effects of spatial attention on the recently discovered ERP signature of evidence accumulation-the central parietal positivity (CPP)-have not been fully characterized. The present study assessed the effect of spatial attention on the C1 and CPP components through a spatially cued contrast discrimination task using stimuli that were specifically designed to produce large-amplitude C1 responses and that varied in sensory evidence strength to characterize the CPP. Participants responded according to which of two checkerboard stimuli had greater contrast following an 80% valid cue toward the upper or lower visual field. Prestimulus alpha power changed topographically based on the cue, suggesting participants shifted attention to prepare for the upcoming stimuli. Despite these attentional shifts in alpha power and the fact that the stimuli reliably elicited C1 responses several times greater than many prior studies, there was no evidence of an attention effect on the C1. The CPP, however, showed a clear increase in build-up rate on valid trials. Our findings suggest that endogenous attention may not affect the early C1 ERP component but may improve behavior at a decision stage, as reflected in brain signals related to evidence accumulation (the CPP).

Behavioral and neural measures of confidence using a novel auditory pitch identification task.

Observers can discriminate between correct versus incorrect perceptual decisions with feelings of confidence. The centro-parietal positivity build-up rate (CPP slope) has been suggested as a likely neural signature of accumulated evidence, which may guide both perceptual performance and confidence. However, CPP slope also covaries with reaction time, which also covaries with confidence in previous studies, and performance and confidence typically covary; thus, CPP slope may index signatures of perceptual performance rather than confidence per se. Moreover, perceptual metacognition-including neural correlates-has largely been studied in vision, with few exceptions. Thus, we lack understanding of domain-general neural signatures of perceptual metacognition outside vision. Here we designed a novel auditory pitch identification task and collected behavior with simultaneous 32-channel EEG in healthy adults. Participants saw two tone labels which varied in tonal distance on each trial (e.g., C vs D, C vs F), then heard a single auditory tone; they identified which label was correct and rated confidence. We found that pitch identification confidence varied with tonal distance, but performance, metacognitive sensitivity (trial-by-trial covariation of confidence with accuracy), and reaction time did not. Interestingly, however, while CPP slope covaried with performance and reaction time, it did not significantly covary with confidence. We interpret these results to mean that CPP slope is likely a signature of first-order perceptual processing and not confidence-specific signals or computations in auditory tasks. Our novel pitch identification task offers a valuable method to examine the neural correlates of auditory and domain-general perceptual confidence.

Neural Signatures of Evidence Accumulation Encode Subjective Perceptual Confidence Independent of Performance.

Confidence is an adaptive computation when environmental feedback is absent, yet there is little consensus regarding how perceptual confidence is computed in the brain. Difficulty arises because confidence correlates with other factors, such as accuracy, response time (RT), or evidence quality. We investigated whether neural signatures of evidence accumulation during a perceptual choice predict subjective confidence independently of these factors. Using motion stimuli, a central-parietal positive-going electroencephalogram component (CPP) behaves as an accumulating decision variable that predicts evidence quality, RT, accuracy, and confidence (Experiment 1, N = 25 adults). When we psychophysically varied confidence while holding accuracy constant (Experiment 2, N = 25 adults), the CPP still predicted confidence. Statistically controlling for RT, accuracy, and evidence quality (Experiment 3, N = 24 adults), the CPP still explained unique variance in confidence. The results indicate that a predecision neural signature of evidence accumulation, the CPP, encodes subjective perceptual confidence in decision-making independent of task performance.

Double Dissociation of Spontaneous Alpha-Band Activity and Pupil-Linked Arousal on Additive and Multiplicative Perceptual Gain.

Perception is a probabilistic process dependent on external stimulus properties and one's internal state. However, which internal states influence perception and via what mechanisms remain debated. We studied how spontaneous alpha-band activity (8-13 Hz) and pupil fluctuations impact visual detection and confidence across stimulus contrast levels (i.e., the contrast response function, CRF). In human subjects of both sexes, we found that low prestimulus alpha power induced an "additive" shift in the CRF, whereby stimuli were reported present more frequently at all contrast levels, including contrast of zero (i.e., false alarms). Conversely, prestimulus pupil size had a "multiplicative" effect on detection such that stimuli occurring during large pupil states (putatively corresponding to higher arousal) were perceived more frequently as contrast increased. Signal detection modeling reveals that alpha power changes detection criteria equally across the CRF but not detection sensitivity (d'), whereas pupil-linked arousal modulated sensitivity, particularly for higher contrasts. Interestingly, pupil size and alpha power were positively correlated, meaning that some of the effect of alpha on detection may be mediated by pupil fluctuations. However, pupil-independent alpha still induced an additive shift in the CRF corresponding to a criterion effect. Our data imply that low alpha boosts detection and confidence by an additive factor, rather than by a multiplicative scaling of contrast responses, a profile which captures the effect of pupil-linked arousal. We suggest that alpha power and arousal fluctuations have dissociable effects on behavior. Alpha reflects the baseline level of visual excitability, which can vary independent of arousal.

Alpha-band Brain Dynamics and Temporal Processing: An Introduction to the Special Focus.

For decades, the intriguing connection between the human alpha rhythm (an 8- to 13-Hz oscillation maximal over posterior cortex) and temporal processes in perception has furnished a rich landscape of proposals. The past decade, however, has seen a surge in interest in the topic, bringing new theoretical, analytic, and methodological developments alongside fresh controversies. This Special Focus on alpha-band dynamics and temporal processing provides an up-to-date snapshot of the playing field, with contributions from leading researchers in the field spanning original perspectives, new evidence, comprehensive reviews and meta-analyses, as well as discussion of ongoing controversies and paths forward. We hope that the perspectives captured here will help catalyze future research and shape the pathways toward a theoretically grounded and mechanistic account of the link between alpha dynamics and temporal properties of perception.

Alpha-Band Frequency and Temporal Windows in Perception: A Review and Living Meta-analysis of 27 Experiments (and Counting).

Temporal windows in perception refer to windows of time within which distinct stimuli interact to influence perception. A simple example is two temporally proximal stimuli fusing into a single percept. It has long been hypothesized that the human alpha rhythm (an 8- to 13-Hz neural oscillation maximal over posterior cortex) is linked to temporal windows, with higher frequencies corresponding to shorter windows and finer-grained temporal resolution. This hypothesis has garnered support from studies demonstrating a correlation between individual differences in alpha-band frequency (IAF) and behavioral measures of temporal processing. However, nonsignificant effects have also been reported. Here, we review and meta-analyze 27 experiments correlating IAF with measures of visual and audiovisual temporal processing. Our results estimate the true correlation in the population to be between .39 and .53, a medium-to-large effect. The effect held when considering visual or audiovisual experiments separately, when examining different IAF estimation protocols (i.e., eyes open and eyes closed), and when using analysis choices that favor a null result. Our review shows that (1) effects have been internally and independently replicated, (2) several positive effects are based on larger sample sizes than the null effects, and (3) many reported null effects are actually in the direction predicted by the hypothesis. A free interactive web app was developed to allow users to replicate our meta-analysis and change or update the study selection at will, making this a "living" meta-analysis (randfxmeta.streamlit.app). We discuss possible factors underlying null reports, design recommendations, and open questions for future research.

Unique effects of sedatives, dissociatives, psychedelics, stimulants, and cannabinoids on episodic memory: A review and reanalysis of acute drug effects on recollection, familiarity, and metamemory.

Despite distinct classes of psychoactive drugs producing putatively unique states of consciousness, there is surprising overlap in terms of their effects on episodic memory and cognition more generally. Episodic memory is supported by multiple subprocesses that have been mostly overlooked in psychopharmacology and could differentiate drug classes. Here, we reanalyzed episodic memory confidence ratings from 10 previously published data sets (28 drug conditions total) using signal detection models to estimate two conscious states involved in episodic memory and one consciously controlled metacognitive process of memory: autonoetic retrieval of specific details (recollection), noetic recognition absent of retrieved details (familiarity), and retrospective introspection of memory decisions (metamemory). Sedatives, dissociatives, psychedelics, stimulants, and cannabinoids had unique patterns of effects on these mnemonic processes dependent on whether they impacted encoding, consolidation, or retrieval (the formation, stabilization, and access to memory traces, respectively). Sedatives at encoding reliably impaired both recollection and familiarity but at consolidation enhanced recollection. Dissociatives and cannabinoids at encoding impaired recollection but less reliably impaired familiarity, and cannabinoids at retrieval increased false recollections. These drug-induced encoding impairments occasionally came with metamemory enhancements, perhaps because of less interstimulus interference. Psychedelics at encoding impaired recollection but tended to enhance familiarity and did not impact metamemory. Stimulants at encoding enhanced metamemory, at consolidation impaired metamemory, and at retrieval enhanced familiarity and metamemory. These findings allude to mechanisms underlying the idiosyncratic phenomena of drugs, such as blackouts from sedatives and presque vu from psychedelics. Finally, these findings converge on a model in which memory quantity and stability influence metamemory. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Task feedback suggests a post-perceptual component to serial dependence.

Decisions across a range of perceptual tasks are biased toward past stimuli. Such serial dependence is thought to be an adaptive low-level mechanism that promotes perceptual stability across time. However, recent studies suggest post-perceptual mechanisms may also contribute to serially biased responses, calling into question a single locus of serial dependence and the nature of integration of past and present sensory inputs. We measured serial dependence in the context of a three-dimensional (3D) motion perception task where uncertainty in the sensory information varied substantially from trial to trial. We found that serial dependence varied with stimulus properties that impact sensory uncertainty on the current trial. Reduced stimulus contrast was associated with an increased bias toward the stimulus direction of the previous trial. Critically, performance feedback, which reduced sensory uncertainty, abolished serial dependence. These results provide clear evidence for a post-perceptual locus of serial dependence in 3D motion perception and support the role of serial dependence as a response strategy in the face of substantial sensory uncertainty.

Brain signatures indexing variation in internal processing during perceptual decision-making.

Brain activity is highly variable during a task. Discovering, characterizing, and linking variability in brain activity to internal processes has primarily relied on experimental manipulations. However, changes in internal processing could arise from many factors independent of experimental conditions. Here we utilize a data-driven clustering method based on modularity-maximation to identify consistent spatial-temporal EEG activity patterns across individual trials. Subjects (N = 25) performed a motion discrimination task with six interleaved levels of coherence. Clustering identified two discrete subtypes of trials with different patterns of activity. Surprisingly, Subtype 1 occurred more frequently in trials with lower motion coherence but was associated with faster response times. Computational modeling suggests that Subtype 1 was characterized by a lower threshold for reaching a decision. These results highlight across-trial variability in decision processes traditionally hidden to experimenters and provide a method for identifying endogenous brain state variability relevant to cognition and behavior.

Evaluating the Evidence for the Functional Inhibition Account of Alpha-band Oscillations during Preparatory Attention.

The functional inhibition account states that alpha-band (8-14 Hz) power implements attentional control by selectively inhibiting task-irrelevant neural representations. This account has been well supported by decades of correlational research showing attention-related changes in the topography of alpha power in anticipation of task-relevant stimuli and is a viable theory of how attention impacts sensory processing, namely, via alpha power changes in sensory areas before stimulus onset. In addition, attention is known to modulate neural responses to stimuli themselves. Thus, a critical prediction of the functional inhibition account is that preparatory alpha modulations should explain variance in the degree of attention-related modulation of neural responses to stimuli. The present article sought evidence for or against this prediction by scouring the literature on attention and alpha oscillations to review papers that explicitly correlated attention-related changes in prestimulus alpha with attention-related changes in stimulus-evoked neural activity. Surprisingly, out of over 100 papers that were examined, we found only nine that explicitly computed such relationships. The results of these nine papers were mixed, with some in support and some arguing against the functional inhibition account of alpha. Our synthesis draws out common design features that may help explain when effects are observed or not. Even among studies that do find correlations, there is inconsistency as to whether preparatory alpha modulations are predictive of sensory or postsensory components of stimulus responses, highlighting avenues for future research. A clear outcome of this review is that future studies on the role of alpha in attentional processing should analyze correlations between attention effects on alpha and attention effects on stimulus-evoked activity, as more data pertinent to this hypothesized relationship are needed.

Individual alpha frequency appears unrelated to the latency of early visual responses.

A large body of work has linked neural oscillations in the alpha-band (8-13 Hz) to visual perceptual outcomes. In particular, studies have found that alpha phase prior to stimulus onset predicts stimulus detection, and sensory responses and that the frequency of alpha can predict temporal properties of perception. These findings have bolstered the idea that alpha-band oscillations reflect rhythmic sampling of visual information, however the mechanisms of this are unclear. Recently two contrasting hypotheses have been proposed. According to the rhythmic perception account, alpha oscillations impose phasic inhibition on perceptual processing and primarily modulate the amplitude or strength of visual responses and thus the likelihood of stimulus detection. On the other hand, the discrete perception account proposes that alpha activity discretizes perceptual inputs thereby reorganizing the timing (not only the strength) of perceptual and neural processes. In this paper, we sought neural evidence for the discrete perception account by assessing the correlation between individual alpha frequencies (IAF) and the latency of early visual evoked event-related potential (ERP) components. If alpha cycles were responsible for shifting neural events in time, then we may expect higher alpha frequencies to be associated with earlier afferent visual ERPs. Participants viewed large checkerboard stimuli presented to either the upper or lower visual field that were designed to elicit a large C1 ERP response (thought to index feedforward primary visual cortex activation). We found no reliable correlation between IAF and the C1 latency, or subsequent ERP component latencies, suggesting that the timing of these visual-evoked potentials was not modulated by alpha frequency. Our results thus fail to find evidence for discrete perception at the level of early visual responses but leave open the possibility of rhythmic perception.

Effects of satisfying and violating expectations on serial dependence.

Serial dependence refers to the phenomenon that observers tend to report stimuli as being more similar to previous stimuli than they really are (attractive dependence) or, in some cases, as more different than they really are (repulsive dependence). Numerous experiments have demonstrated serial dependence for a range of modalities and stimulus features, highlighting the role of bottom-up sensory interactions. However, comparatively less research has focused on how higher-level cognitive factors, such as expectations, might influence serial dependence. Here, we manipulated expectations by having observers respond to target luminance gratings that occurred at the end of a sequence of non-target gratings. The sequence either rotated predictably (inducing an expectation), varied randomly (inducing no expectation), or rotated predictably but had a random target orientation (violating expectations). We found that observers produced less errors and indicated less uncertainty in their estimations of expected stimuli but their responses were biased away from the penultimate stimulus in the sequence (repulsive dependence). In contrast, following random sequences, responses showed an attractive bias to the penultimate stimulus in the sequence. Unexpected targets showed a mixture of both biases, such that when targets happened (by chance) to appear as expected, responses were repulsed, but responses to target orientations that more clearly violated expectations were attracted. These results indicate that, whereas attraction to previous stimuli may be a default strategy employed in response to random and unexpected events, certain expectations can reverse the default bias into a repulsive one.

Brain signatures indexing variation in internal processing during perceptual decision-making.

Brain activity is highly variable even while performing the same cognitive task with consequences for performance. Discovering, characterizing, and linking variability in brain activity to internal processes has primarily relied on experimentally inducing changes (e.g., via attention manipulation) to identify neuronal and behavioral consequences or studying spontaneous changes in ongoing brain dynamics. However, changes in internal processing could arise from many factors, such as variation in strategy or arousal, that are independent of experimental conditions. Here we utilize a data-driven clustering method based on modularity-maximation to identify consistent spatial-temporal EEG activity patterns across individual trials and relate this activity to behavioral performance. Subjects (N = 25) performed a motion direction discrimination task with six interleaved levels of motion coherence. Modularity-maximization based clustering identified two discrete spatial-temporal clusters, or subtypes, of trials with different patterns of brain activity. Surprisingly, even though Subtype 1 occurred more frequently with lower motion coherence, it was nonetheless associated with faster response times. Computational modeling suggests that Subtype 1 was characterized by a lower threshold for reaching a decision. These results highlight trial-to-trial variability in decision processes usually masked to experimenters and provide a method for identifying endogenous brain state variability relevant to cognition and behavior.

Context Binding in Visual Working Memory Is Reflected in Bilateral Event-Related Potentials, But Not in Contralateral Delay Activity.

Successful retrieval of a specific item from visual working memory (VWM) depends on the binding of that item to its unique context. Recent functional magnetic resonance imaging studies of VWM manipulating memory set homogeneity have identified an important role for the intraparietal sulcus in context binding, independent of any role in representing stimulus identity. The current study explored whether the contralateral delay activity (CDA), which is an event-related potential (ERP) component derived from posterior electrodes that tracks the amount of information held in VWM, might also be sensitive to context-binding demands. In experiment 1, human participants performed lateralized delayed recognition with memory sets containing one, three, or five items that were drawn from the same category (orientations: "homogeneous") or from different categories (orientation, color, and luminance: "heterogeneous"). Because the location and identity of the memory probe indicated the item to be retrieved, homogeneous trials placed higher context-binding demands. VWM capacity was higher in heterogeneous trials. ERPs contralateral (contra) and ipsilateral (ipsi) to the remembered stimuli were higher for homogeneous trials, but these differences were removed in the contra - ipsi subtraction that produced the CDA. In experiment 2, human participants performed lateralized delayed recall with memory sets of one or three items (homogeneous or heterogeneous). Behavior was superior for three-item heterogeneous trials than for homogeneous trials, with modeling revealing context-binding errors in the latter. Bilateral ERPs and CDA results replicated experiment 1. These results support that the CDA tracks the number of object files engaged by VWM and establish that it is not sensitive to context-binding demands.

Consensus Goals in the Field of Visual Metacognition.

Despite the tangible progress in psychological and cognitive sciences over the last several years, these disciplines still trail other more mature sciences in identifying the most important questions that need to be solved. Reaching such consensus could lead to greater synergy across different laboratories, faster progress, and increased focus on solving important problems rather than pursuing isolated, niche efforts. Here, 26 researchers from the field of visual metacognition reached consensus on four long-term and two medium-term common goals. We describe the process that we followed, the goals themselves, and our plans for accomplishing these goals. If this effort proves successful within the next few years, such consensus building around common goals could be adopted more widely in psychological science.

The positive evidence bias in perceptual confidence is unlikely post-decisional.

Confidence in a perceptual decision is a subjective estimate of the accuracy of one's choice. As such, confidence is thought to be an important computation for a variety of cognitive and perceptual processes, and it features heavily in theorizing about conscious access to perceptual states. Recent experiments have revealed a "positive evidence bias" (PEB) in the computations underlying confidence reports. A PEB occurs when confidence, unlike objective choice, overweights the evidence for the correct (or chosen) option, relative to evidence against the correct (or chosen) option. Accordingly, in a perceptual task, appropriate stimulus conditions can be arranged that produce selective changes in confidence reports but no changes in accuracy. Although the PEB is generally assumed to reflect the observer's perceptual and/or decision processes, post-decisional accounts have not been ruled out. We therefore asked whether the PEB persisted under novel conditions that addressed two possible post-decisional accounts: (i) post-decision evidence accumulation that contributes to a confidence report solicited after the perceptual choice and (ii) a memory bias that emerges in the delay between the stimulus offset and the confidence report. We found that even when the stimulus remained on the screen until observers responded, and when observers reported their choice and confidence simultaneously, the PEB still emerged. Signal detection-based modeling showed that the PEB was not associated with changes to metacognitive efficiency, but rather to confidence criteria. The data show that memory biases cannot explain the PEB and provide evidence against a post-decision evidence accumulation account, bolstering the idea that the PEB is perceptual or decisional in nature.

Spontaneous alpha-band amplitude predicts subjective visibility but not discrimination accuracy during high-level perception.

Near-threshold perception is a paradigm case of awareness diverging from reality - the perception of an unchanging stimulus can vacillate from undetected to clearly perceived. The amplitude of low-frequency brain oscillations - particularly in the alpha-band (8-13 Hz) - has emerged as a reliable predictor of trial-to-trial variability in perceptual decisions based on simple, low-level stimuli. Here, we addressed the question of how spontaneous oscillatory amplitude impacts subjective and objective aspects of perception using high-level visual stimuli. Human observers completed a near-threshold face/house discrimination task with subjective visibility ratings while electroencephalograms (EEG) were recorded. Using single-trial multiple regression analysis, we found that spontaneous fluctuations in prestimulus alpha-band amplitude were negatively related to visibility judgments but did not predict trial-by-trial accuracy. These results extend previous findings that indicate that strong prestimulus alpha diminishes subjective perception without affecting the accuracy or sensitivity (d') of perceptual decisions into the domain of high-level perception.

Spectral Distribution Dynamics across Different Attentional Priority States.

Anticipatory covert spatial attention improves performance on tests of visual detection and discrimination, and shifts are accompanied by decreases and increases of α band power at electroencephalography (EEG) electrodes corresponding to the attended and unattended location, respectively. Although the increase at the unattended location is often interpreted as an active mechanism (e.g., inhibiting processing at the unattended location), most experiments cannot rule out the alternative possibility that it is a secondary consequence of selection elsewhere. To adjudicate between these accounts, we designed a Posner-style visual cueing task in which male and female human participants made orientation judgments of targets appearing at one of four locations: up, down, right, or left. Critically, trials were blocked such that within a block the locations along one meridian alternated in status between attended and unattended, and targets never appeared at the other two, making them irrelevant. Analyses of the concurrently measured EEG signal were conducted on "traditional" narrowband α (8-14 Hz), as well as on two components resulting from the decomposition of this signal: "periodic" α; and the slope of the aperiodic 1/f-like component. Although data from right-left blocks replicated the familiar pattern of lateralized asymmetry in narrowband α power, with neither α signal could we find evidence for any difference in the time course at unattended versus irrelevant locations, an outcome consistent with the secondary-consequence interpretation of attention-related dynamics in the α band. Additionally, 1/f slope was shallower at attended and unattended locations, relative to irrelevant, suggesting a tonic adjustment of physiological state.SIGNIFICANCE STATEMENT Visual spatial attention, the prioritization of one location in the visual field, is critical for guiding behavior in cluttered environments. Although influential theories posit an important role for α band oscillations in the inhibition of processing at unattended locations, we used a novel procedure to find evidence for an alternative interpretation: selection of one location may simply result in a return to physiological baseline at all others. In addition to determining one way that attention does not work (important for future progress in this field), we also discovered novel evidence for one way that it does work: by modifying the tonic physiological state (indexed by an aperiodic component of the electroencephalography (EEG)] at locations where spatial selection is likely to occur.

Pre-stimulus alpha-band phase gates early visual cortex responses.

Alpha-band (8-13 Hz) oscillations have been shown to phasically inhibit perceptual reports in human observers, yet the underlying physiological mechanism of this effect is debated. According to contrasting models, based primarily on animal experiments, alpha activity is thought to either originate from specialized cells in the visual thalamus and periodically inhibit the relay of visual information to the primary visual cortex (V1) in a feedforward manner, or to propagate from higher visual areas back to V1 in a feedback manner. Human neurophysiological evidence in favor of either hypothesis, both, or neither, has been limited. To help address this issue, we explored the link between pre-stimulus alpha phase and visual electroencephalography (EEG) responses thought to arise from afferent input onto human V1. Specially-designed visual stimuli were used to elicit large amplitude C1 event-related potentials (ERP), with polarity, topography, and timing indicative of striate genesis. Single-trial circular-linear associations between pre-stimulus phase and post-stimulus global field power (GFP) during the C1 time window revealed significant effects peaking in the alpha frequency band. Control analyses ruling out the potential confound of post-stimulus data bleeding into the pre-stimulus window demonstrated that GFP amplitude decreases as pre-stimulus alpha phase deviates from an individual's preferred phase. These findings demonstrate an early locus - suggesting that the phase of pre-stimulus alpha oscillations could modulate visual processing by gating the feedforward flow of sensory input between the thalamus and V1, although other models are potentially compatible.