Sustained attention operates via dissociable neural mechanisms across different eccentric locations.

In primates, foveal and peripheral vision have distinct neural architectures and functions. However, it has been debated if selective attention operates via the same or different neural mechanisms across eccentricities. We tested these alternative accounts by examining the effects of selective attention on the steady-state visually evoked potential (SSVEP) and the fronto-parietal signal measured via EEG from human subjects performing a sustained visuospatial attention task. With a negligible level of eye movements, both SSVEP and SND exhibited the heterogeneous patterns of attentional modulations across eccentricities. Specifically, the attentional modulations of these signals peaked at the parafoveal locations and such modulations wore off as visual stimuli appeared closer to the fovea or further away towards the periphery. However, with a relatively higher level of eye movements, the heterogeneous patterns of attentional modulations of these neural signals were less robust. These data demonstrate that the top-down influence of covert visuospatial attention on early sensory processing in human cortex depends on eccentricity and the level of saccadic responses. Taken together, the results suggest that sustained visuospatial attention operates differently across different eccentric locations, providing new understanding of how attention augments sensory representations regardless of where the attended stimulus appears.

Feedforward attentional selection in sensory cortex.

Salient objects grab attention because they stand out from their surroundings. Whether this phenomenon is accomplished by bottom-up sensory processing or requires top-down guidance is debated. We tested these alternative hypotheses by measuring how early and in which cortical layer(s) neural spiking distinguished a target from a distractor. We measured synaptic and spiking activity across cortical columns in mid-level area V4 of male macaque monkeys performing visual search for a color singleton. A neural signature of attentional capture was observed in the earliest response in the input layer 4. The magnitude of this response predicted response time and accuracy. Errant behavior followed errant selection. Because this response preceded top-down influences and arose in the cortical layer not targeted by top-down connections, these findings demonstrate that feedforward activation of sensory cortex can underlie attentional priority.

Dissociable Neural Mechanisms Underlie the Effects of Attention on Visual Appearance and Response Bias.

A prominent theoretical framework spanning philosophy, psychology, and neuroscience holds that selective attention penetrates early stages of perceptual processing to alter the subjective visual experience of behaviorally relevant stimuli. For example, searching for a red apple at the grocery store might make the relevant color appear brighter and more saturated compared with seeing the exact same red apple while searching for a yellow banana. In contrast, recent proposals argue that data supporting attention-related changes in appearance reflect decision- and motor-level response biases without concurrent changes in perceptual experience. Here, we tested these accounts by evaluating attentional modulations of EEG responses recorded from male and female human subjects while they compared the perceived contrast of attended and unattended visual stimuli rendered at different levels of physical contrast. We found that attention enhanced the amplitude of the P1 component, an early evoked potential measured over visual cortex. A linking model based on signal detection theory suggests that response gain modulations of the P1 component track attention-induced changes in perceived contrast as measured with behavior. In contrast, attentional cues induced changes in the baseline amplitude of posterior alpha band oscillations (∼9-12 Hz), an effect that best accounts for cue-induced response biases, particularly when no stimuli are presented or when competing stimuli are similar and decisional uncertainty is high. The observation of dissociable neural markers that are linked to changes in subjective appearance and response bias supports a more unified theoretical account and demonstrates an approach to isolate subjective aspects of selective information processing.SIGNIFICANCE STATEMENT Does attention alter visual appearance, or does it simply induce response bias? In the present study, we examined these competing accounts using EEG and linking models based on signal detection theory. We found that response gain modulations of the visually evoked P1 component best accounted for attention-induced changes in visual appearance. In contrast, cue-induced baseline shifts in alpha band activity better explained response biases. Together, these results suggest that attention concurrently impacts visual appearance and response bias, and that these processes can be experimentally isolated.

Statistical learning speeds visual search: More efficient selection, or faster response?

Learning statistical regularities of target objects speeds visual search performance. However, we do not yet know whether this statistical learning effect is driven by biasing attentional selection at the early perceptual stage of processing, as theories of attention propose, or by improving the decision-making efficiency at a late response-related stage. Leveraging the high-temporal resolution of the event-related potential (ERP) technique, we had 16 human observers perform a visual search task where we inserted a fine-grained statistical regularity that the target shapes appeared in different colors with six unique probabilities. Observers unintentionally learned these regularities such that they were faster to report targets that appeared in more likely target colors. The observers' ERPs showed that this learning effect resulted in subjects making faster decisions about the target presence, and not by preferentially shifting attention to more rapidly select likely target colors, as is often assumed by the attentional selection account, supporting a post-selection account for statistical learning of the probabilistic regularities of target features. These results provide fundamental insights into the attentional control mechanisms of statistical learning. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Cross-frequency coupling of frontal theta and posterior alpha is unrelated to the fidelity of visual long-term memory encoding.

Given that the encoding of information into visual long-term memory relies on multiple spatially distinct areas in the human brain, encoding into visual memory is a cognitive process likely to rely on networks formed via large-scale coupled neuronal oscillations. Previous research suggests that decreases in occipital alpha power and increases in mid-frontal theta power individually contribute to the encoding of retrievable representations in visual long-term memory. The present study asks whether these oscillations form a coupled network that operates during long-term memory encoding. Here we show that neither amplitude-amplitude coupling nor phase-amplitude coupling between frontal theta and posterior alpha are correlated with observers' subsequent memory when storing pictures in visual long-term memory. Correlations between alpha and theta power were stronger during eyes-open, resting-state periods when no task was performed than immediately following the presentation of a to-be-remembered picture. Finally, we also found that the strength of theta-alpha coupling was not modified by temporal lobe anodal transcranial direct current stimulation, despite observers showing enhanced memory for pictures behaviorally. Collectively, these findings indicate that posterior alpha and frontal theta activity are not simply component parts of a larger scale coupled network underlying visual memory.

Does motor noise contaminate estimates of the precision of visual working memory?

The continuous-report task, in which subjects report the color of visual working memory representation by clicking on a color wheel, has become the gold standard for measuring the precision and number of representations stored in visual working memory. This task requires fine motor control, typically with a mouse, but the precision of responses have been interpreted as being entirely due to the precision of the memory representations, without regard to the contribution of noise from the response effectors (i.e., motor control of the hand). Here we tested the seemingly likely possibility that motor noise contaminates our estimates of visual memory representations in the continuous-report task by simply asking subjects to complete the color wheel continuous-report task using either their dominant or non-dominant hand on different blocks of trials. We found that subjects took longer to complete the task with their non-dominant hand, but this did not affect the precision of their responses. Our findings suggest that this commonly used task to study visual memory may be relatively immune from contamination by motor noise at the output stage.

Resolving the mesoscopic missing link: Biophysical modeling of EEG from cortical columns in primates.

Event-related potentials (ERP) are among the most widely measured indices for studying human cognition. While their timing and magnitude provide valuable insights, their usefulness is limited by our understanding of their neural generators at the circuit level. Inverse source localization offers insights into such generators, but their solutions are not unique. To address this problem, scientists have assumed the source space generating such signals comprises a set of discrete equivalent current dipoles, representing the activity of small cortical regions. Based on this notion, theoretical studies have employed forward modeling of scalp potentials to understand how changes in circuit-level dynamics translate into macroscopic ERPs. However, experimental validation is lacking because it requires in vivo measurements of intracranial brain sources. Laminar local field potentials (LFP) offer a mechanism for estimating intracranial current sources. Yet, a theoretical link between LFPs and intracranial brain sources is missing. Here, we present a forward modeling approach for estimating mesoscopic intracranial brain sources from LFPs and predict their contribution to macroscopic ERPs. We evaluate the accuracy of this LFP-based representation of brain sources utilizing synthetic laminar neurophysiological measurements and then demonstrate the power of the approach in vivo to clarify the source of a representative cognitive ERP component. To that end, LFP was measured across the cortical layers of visual area V4 in macaque monkeys performing an attention demanding task. We show that area V4 generates dipoles through layer-specific transsynaptic currents that biophysically recapitulate the ERP component through the detailed forward modeling. The constraints imposed on EEG production by this method also revealed an important dissociation between computational and biophysical contributors. As such, this approach represents an important bridge between laminar microcircuitry, through the mesoscopic activity of cortical columns to the patterns of EEG we measure at the scalp.

Even affective changes induced by the global health crisis are insufficient to perturb the hyper-stability of visual long-term memory.

Past studies of emotion and mood on memory have mostly focused on the learning of emotional material in the laboratory or on the consequences of a punctate catastrophic event. However, the influence of a long-lasting global condition on memory and learning has not been studied. The COVID-19 pandemic unfortunately offered a unique situation to observe the effects of prolonged, negative events on human memory for visual information. One thousand online subjects were asked to remember the details of real-world photographs of objects to enable fine-grained visual discriminations from novel within-category foils. Visual memory performance was invariant across time, regardless of the infection rate in the local or national population, or the subjects' self-reported affective state using the Positive and Negative Affect Schedule (PANAS). Thus, visual memory provides the human brain with storage that is particularly resilient to changes in emotional state, even when those changes are experienced for months longer than any imaginable laboratory procedure.

A meta-analytic review of transcranial direct current stimulation (tDCS) on general psychopathology symptoms of schizophrenia; immediate improvement followed by a return to baseline.

Transcranial direct current stimulation (tDCS) is a promising tool for alleviating positive and negative symptoms of schizophrenia, but its role in functional outcome remains uncertain. This meta-analysis examined the effects of tDCS on general psychopathology symptoms (GPS) from the Positive and Negative Syndrome Scale (PANSS) because GPS are closely associated with daily functioning. Literature search using Medline and PsycINFO identified 8 randomized controlled trials with tDCS and PANSS. The GPS were significantly reduced after tDCS but there was no evidence for long-term treatment effects. Further research is needed to optimize the dosing of tDCS and to understand individual differences in treatment response.

Laminar microcircuitry of visual cortex producing attention-associated electric fields.

Cognitive operations are widely studied by measuring electric fields through EEG and ECoG. However, despite their widespread use, the neural circuitry giving rise to these signals remains unknown because the functional architecture of cortical columns producing attention-associated electric fields has not been explored. Here, we detail the laminar cortical circuitry underlying an attention-associated electric field measured over posterior regions of the brain in humans and monkeys. First, we identified visual cortical area V4 as one plausible contributor to this attention-associated electric field through inverse modeling of cranial EEG in macaque monkeys performing a visual attention task. Next, we performed laminar neurophysiological recordings on the prelunate gyrus and identified the electric-field-producing dipoles as synaptic activity in distinct cortical layers of area V4. Specifically, activation in the extragranular layers of cortex resulted in the generation of the attention-associated dipole. Feature selectivity of a given cortical column determined the overall contribution to this electric field. Columns selective for the attended feature contributed more to the electric field than columns selective for a different feature. Last, the laminar profile of synaptic activity generated by V4 was sufficient to produce an attention-associated signal measurable outside of the column. These findings suggest that the top-down recipient cortical layers produce an attention-associated electric field that can be measured extracortically with the relative contribution of each column depending upon the underlying functional architecture.

Alpha suppression indexes a spotlight of visual-spatial attention that can shine on both perceptual and memory representations.

Although researchers have been recording the human electroencephalogram (EEG) for almost a century, we still do not completely understand what cognitive processes are measured by the activity of different frequency bands. The 8- to 12-Hz activity in the alpha band has long been a focus of this research, but our understanding of its links to cognitive mechanisms has been rapidly evolving recently. Here, we review and discuss the existing evidence for two competing perspectives about alpha activity. One view proposes that the suppression of alpha-band power following the onset of a stimulus array measures attentional selection. The competing view is that this same activity measures the buffering of the task-relevant representations in working memory. We conclude that alpha-band activity following the presentation of stimuli appears to be due to the operation of an attentional selection mechanism, with characteristics that mirror the classic views of attention as selecting both perceptual inputs and representations already stored in memory.

Induced Forgetting Is the Result of True Forgetting, Not Shifts in Decision-making Thresholds.

Induced forgetting occurs when accessing an item in memory appears to harm memory representations of categorically related items. However, it is possible that the actual memory representations are unharmed. Instead, people may just change how they make decisions. Specifically, signal detection theory suggests this apparent forgetting may be due to participants shifting their decision criterion. Here, we used behavioral and electrophysiological measures to determine whether induced forgetting is truly due to changes in how items are represented or simply due to a shifting criterion. Participants' behavior and brain activity showed that induced forgetting was due to changes in the strength of the underlying representations, weighing against a criterion shift explanation of induced forgetting.

Medium strength visual long-term memories are the most fragile.

What memories do humans forget? One theory proposes that memories stored with moderate activation levels are weakened when faced with competitive stress so that they are particularly prone to be forgotten. However, research suggests that visual long-term memories are stronger than memories of other modalities, and therefore may never fall into this moderate activation zone. Here we tested these competing predictions by showing to-be-remembered pictures while we recorded event-related potentials (ERPs) indexing memory activation during encoding. We found that visual memories with medium levels of activation when first encoded were more prone to forgetting than memories with high or low encoding activation levels, but this only occurred if a memory was faced with competition. This study shows that we forget moderately activated memories when they are subjected to competition, regardless of the modality of experience.

Spatial location is filtered out of visual working memory representations when task irrelevant, just like other features.

Contrary to early proposals stating that we always store bound object features in visual working memory, more recent work has suggested that we can inhibit the encoding, or consolidation, of irrelevant features of objects into visual working memory. However, a number of theoretical proposals suggest that spatial location is a special feature of an object that might be obligatorily bound to objects stored in visual working memory. In this study, I used a masking paradigm to measure the efficiency of encoding into visual working memory while subjects were tasked with remembering the location, color, or both of these features of the objects. The measures of consolidation efficiency indicate that spatial location is not encoded into visual working memory unless it is relevant for the task at hand. Thus, the present experiments show that we can control which features of an object are selectively stored in working memory, including spatial location, a feature thought to be immune to such filtering.

α-Band activity tracks a two-dimensional spotlight of attention during spatial working memory maintenance.

Covert spatial attention is thought to facilitate the maintenance of locations in working memory, and EEG α-band activity (8-12 Hz) is proposed to track the focus of covert attention. Recent work has shown that multivariate patterns of α-band activity track the polar angle of remembered locations relative to fixation. However, a defining feature of covert spatial attention is that it facilitates processing in a specific region of the visual field, and prior work has not determined whether patterns of α-band activity track the two-dimensional (2-D) coordinates of remembered stimuli within a visual hemifield or are instead maximally sensitive to the polar angle of remembered locations around fixation. Here, we used a lateralized spatial estimation task, in which observers remembered the location of one or two target dots presented to one side of fixation, to test this question. By applying a linear discriminant classifier to the topography of α-band activity, we found that we were able to decode the location of remembered stimuli. Critically, model comparison revealed that the pattern of classifier choices observed across remembered positions was best explained by a model assuming that α-band activity tracks the 2-D coordinates of remembered locations rather than a model assuming that α-band activity tracks the polar angle of remembered locations relative to fixation. These results support the hypothesis that this α-band activity is involved in the spotlight of attention, and arises from mid- to lower-level visual areas involved in maintaining spatial locations in working memory.NEW & NOTEWORTHY A substantial body of work has shown that patterns of EEG α-band activity track the angular coordinates of attended and remembered stimuli around fixation, but whether these patterns track the two-dimensional coordinates of stimuli presented within a visual hemifield remains an open question. Here, we demonstrate that α-band activity tracks the two-dimensional coordinates of remembered stimuli within a hemifield, showing that α-band activity reflects a spotlight of attention focused on locations maintained in working memory.

Stimulus-induced Alpha Suppression Tracks the Difficulty of Attentional Selection, Not Visual Working Memory Storage.

Human alpha-band activity (8-12 Hz) has been proposed to index a variety of mechanisms during visual processing. Here, we distinguished between an account in which alpha suppression indexes selective attention versus an account in which it indexes subsequent working memory storage. We manipulated two aspects of the visual stimuli that perceptual attention is believed to mitigate before working memory storage: the potential interference from distractors and the size of the focus of attention. We found that the magnitude of alpha-band suppression tracked both of these aspects of the visual arrays. Thus, alpha-band activity after stimulus onset is clearly related to how the visual system deploys perceptual attention and appears to be distinct from mechanisms that store target representations in working memory.

Converging Evidence That Neural Plasticity Underlies Transcranial Direct-Current Stimulation.

It is not definitely known how direct-current stimulation causes its long-lasting effects. Here, we tested the hypothesis that the long time course of transcranial direct-current stimulation (tDCS) is because of the electrical field increasing the plasticity of the brain tissue. If this is the case, then we should see tDCS effects when humans need to encode information into long-term memory, but not at other times. We tested this hypothesis by delivering tDCS to the ventral visual stream of human participants during different tasks (i.e., recognition memory vs. visual search) and at different times during a memory task. We found that tDCS improved memory encoding, and the neural correlates thereof, but not retrieval. We also found that tDCS did not change the efficiency of information processing during visual search for a certain target object, a task that does not require the formation of new connections in the brain but instead relies on attention and object recognition mechanisms. Thus, our findings support the hypothesis that direct-current stimulation modulates brain activity by changing the underlying plasticity of the tissue.

A Minimal Biophysical Model of Neocortical Pyramidal Cells: Implications for Frontal Cortex Microcircuitry and Field Potential Generation.

Ca2+ spikes initiated in the distal trunk of layer 5 pyramidal cells (PCs) underlie nonlinear dynamic changes in the gain of cellular response, critical for top-down control of cortical processing. Detailed models with many compartments and dozens of ionic channels can account for this Ca2+ spike-dependent gain and associated critical frequency. However, current models do not account for all known Ca2+-dependent features. Previous attempts to include more features have required increasing complexity, limiting their interpretability and utility for studying large population dynamics. We overcome these limitations in a minimal two-compartment biophysical model. In our model, a basal-dendrites/somatic compartment included fast-inactivating Na+ and delayed-rectifier K+ conductances, while an apical-dendrites/trunk compartment included persistent Na+, hyperpolarization-activated cation (I h ), slow-inactivating K+, muscarinic K+, and Ca2+ L-type. The model replicated the Ca2+ spike morphology and its critical frequency plus three other defining features of layer 5 PC synaptic integration: linear frequency-current relationships, back-propagation-activated Ca2+ spike firing, and a shift in the critical frequency by blocking I h Simulating 1000 synchronized layer 5 PCs, we reproduced the current source density patterns evoked by Ca2+ spikes and describe resulting medial-frontal EEG on a male macaque monkey. We reproduced changes in the current source density when I h was blocked. Thus, a two-compartment model with five crucial ionic currents in the apical dendrites reproduces all features of these neurons. We discuss the utility of this minimal model to study the microcircuitry of agranular areas of the frontal lobe involved in cognitive control and responsible for event-related potentials, such as the error-related negativity.SIGNIFICANCE STATEMENT A minimal model of layer 5 pyramidal cells replicates all known features crucial for distal synaptic integration in these neurons. By redistributing voltage-gated and returning transmembrane currents in the model, we establish a theoretical framework for the investigation of cortical microcircuit contribution to intracranial local field potentials and EEG. This tractable model will enable biophysical evaluation of multiscale electrophysiological signatures and computational investigation of cortical processing.

Dissociation of Medial Frontal ß-Bursts and Executive Control.

The neural mechanisms of executive and motor control concern both basic researchers and clinicians. In human studies, preparation and cancellation of movements are accompanied by changes in the ß-frequency band (15-29 Hz) of electroencephalogram (EEG). Previous studies with human participants performing stop signal (countermanding) tasks have described reduced frequency of transient ß-bursts over sensorimotor cortical areas before movement initiation and increased ß-bursting over medial frontal areas with movement cancellation. This modulation has been interpreted as contributing to the trial-by-trial control of behavior. We performed identical analyses of EEG recorded over the frontal lobe of macaque monkeys (one male, one female) performing a saccade countermanding task. While we replicate the occurrence and modulation of ß-bursts associated with initiation and cancellation of saccades, we found that ß-bursts occur too infrequently to account for the observed stopping behavior. We also found ß-bursts were more common after errors, but their incidence was unrelated to response time (RT) adaptation. These results demonstrate the homology of this EEG signature between humans and macaques but raise questions about the current interpretation of ß band functional significance.SIGNIFICANCE STATEMENT The finding of increased ß-bursting over medial frontal cortex with movement cancellation in humans is difficult to reconcile with the finding of modulation too late to contribute to movement cancellation in medial frontal cortex of macaque monkeys. To obtain comparable measurement scales, we recorded electroencephalogram (EEG) over medial frontal cortex of macaques performing a stop signal (countermanding) task. We replicated the occurrence and modulation of ß-bursts associated with the cancellation of movements, but we found that ß-bursts occur too infrequently to account for observed stopping behavior. Unfortunately, this finding raises doubts whether ß-bursts can be a causal mechanism of response inhibition, which impacts future applications in devices such as brain-machine interfaces.

What not to look for: Electrophysiological evidence that searchers prefer positive templates.

To-be-attended information can be specified either with positive cues (I'll be wearing a blue shirt) or with negative cues (I won't be wearing a red shirt). Numerous experiments have found that positive cues help search more than negative cues. Given that negative cues produce smaller benefits compared to positive cues, it stands to reason that searchers may choose to use positive templates instead of negative templates if given the opportunity. Here, we evaluate this possibility with behavioral measures as well as by directly measuring the formation of positive and negative templates with event-related potentials. Analysis of the contralateral delay activity (CDA) elicited by cues revealed that positive and negative templates relied on working memory to the same extent, even when negative working memory templates could have been circumvented by relying on long-term memories of target colors. Whereas the CDA did not discriminate positive and negative templates, a CNV-like potential did, suggesting cognitive differences between positive and negative templates beyond visual working memory. However, when both positive and negative information were presented in each cue, participants preferred to make use of the positive cues, as indicated by a CDA contralateral to the positive color in negative cue blocks, and a lack of search benefits for positive- and negative-color cues relative to positive-color cues alone. Our results show that searchers elect to selectively encode only positive information into visual working memory when both positive and negative information are available.