Category Learning Selectively Enhances Representations of Boundary-Adjacent Exemplars in Early Visual Cortex.

Category learning and visual perception are fundamentally interactive processes, such that successful categorization often depends on the ability to make fine visual discriminations between stimuli that vary on continuously valued dimensions. Research suggests that category learning can improve perceptual discrimination along the stimulus dimensions that predict category membership and that these perceptual enhancements are a byproduct of functional plasticity in the visual system. However, the precise mechanisms underlying learning-dependent sensory modulation in categorization are not well understood. We hypothesized that category learning leads to a representational sharpening of underlying sensory populations tuned to values at or near the category boundary. Furthermore, such sharpening should occur largely during active learning of new categories. These hypotheses were tested using fMRI and a theoretically constrained model of vision to quantify changes in the shape of orientation representations while human adult subjects learned to categorize physically identical stimuli based on either an orientation rule (N = 12) or an orthogonal spatial frequency rule (N = 13). Consistent with our predictions, modeling results revealed relatively enhanced reconstructed representations of stimulus orientation in visual cortex (V1-V3) only for orientation rule learners. Moreover, these reconstructed representations varied as a function of distance from the category boundary, such that representations for challenging stimuli near the boundary were significantly sharper than those for stimuli at the category centers. These results support an efficient model of plasticity wherein only the sensory populations tuned to the most behaviorally relevant regions of feature space are enhanced during category learning.

Post-error adjustments occur in both reaching and grasping.

Post-error slowing (PES), the tendency to slow down a behavioral response after a previous error, has typically been investigated during simple cognitive tasks using response time as a measure of PES magnitude. More recently, PES was investigated during a single reach-to-grasp task to determine where post-error adjustments are employed in a more ecological setting. Kinematic analyses in the previous study detected PES during pre-movement planning and within the grasping component of movement execution. In the current study (N = 22), we increased the cognitive demands of a reach-to-grasp task by adding a choice between target and distractor locations to further explore PES, and other post-error adjustments, under different task conditions. We observed a significant main effect of task condition on overall reaction time (RT); however, it did not significantly impact PES or other post-error adjustments. Nonetheless, the results of this study suggest post-error adjustment is a flexible process that can be observed during pre-movement planning and within the onset and magnitude of the reaching component, as well as in the magnitudes of the grasping component. Considering the sum of the results in the context of existing literature, we conclude that the findings add support to a functional account of error reactivity, such that post-error adjustments are implemented intentionally to improve performance.

Changes in pupil size track self-control failure.

People are more likely to perform poorly on a self-control task following a previous task requiring self-control (ego-depletion), but the mechanism for this effect remains unclear. We used pupillometry to test the role of attentional effort in ego-depletion. We hypothesized that an elevated pupil diameter (PD)-a common physiological measure of effort-during an initial task requiring self-control should be negatively associated with performance on a subsequent control task. To test this hypothesis, participants were first assigned to either a high- or low-demand attention task (manipulation; a standard ego-depletion paradigm), after which all participants completed the same Stroop task. We then separately extracted both sustained (low-frequency) and phasic (high-frequency) changes in PD from both tasks to evaluate possible associations with lapses of cognitive control on the Stroop task. We first show that in the initial task, sustained PD was larger among participants who were assigned to the demanding attention condition. Furthermore, ego-depletion effects were serially mediated by PD: an elevated PD response emerged rapidly among the experimental group during the manipulation, persisted as an elevated baseline response during the Stroop task, and predicted worse accuracy on incongruent trials, revealing a potential indirect pathway to ego-depletion via sustained attention. Secondary analyses revealed another, independent and direct pathway via high levels of transient attentional control: participants who exhibited large phasic responses during the manipulation tended to perform worse on the subsequent Stroop task. We conclude by exploring the neuroscientific implications of these results within the context of current theories of self-control.

Space- and feature-based attention operate both independently and interactively within latent components of perceptual decision making.

Top-down visual attention filters undesired stimuli while selected information is afforded the lion's share of limited cognitive resources. Multiple selection mechanisms can be deployed simultaneously, but how unique influences of each combine to facilitate behavior remains unclear. Previously, we failed to observe an additive perceptual benefit when both space-based attention (SBA) and feature-based attention (FBA) were cued in a sparse display (Liang & Scolari, 2020): FBA was restricted to higher order decision-making processes when combined with a valid spatial cue, whereas SBA additionally facilitated target enhancement. Here, we introduced a series of design modifications across three experiments to elicit both attention mechanisms within signal enhancement while also investigating the impacts on decision making. First, we found that when highly reliable spatial and feature cues made unique contributions to search (experiment 1), or when each cue component was moderately reliable (experiments 2a and 2b), both mechanisms were deployed independently to resolve the target. However, the same manipulations produced interactive attention effects within other latent decision-making components that depended on the probability of the integrated cueing object. Time spent before evidence accumulation was reduced and responses were more conservative for the most likely pre-cue combination-even when it included an invalid component. These data indicate that selection mechanisms operate on sensory signals invariably in an independent manner, whereas a higher-order dependency occurs outside of signal enhancement.

Limited interactions between space- and feature-based attention in visually sparse displays.

Top-down visual attention selectively filters sensory input so relevant information receives preferential processing. Feature-based attention (FBA) enhances the representation of relevant low-level features, whereas space-based attention (SBA) enhances information at relevant location(s). The present study investigates whether the unique influences of SBA and FBA combine to facilitate behavior in a perceptually demanding discrimination task. We first demonstrated that, independently, both color and location pre-cues could effectively direct attention to facilitate perceptual decision making of a target. We then examined the combined effects of SBA and FBA in the same design by deploying a predictive color arrow pre-cue. Only SBA effects were observed in performance accuracy and reaction time. However, we detected a reaction time cost when a valid spatial cue was paired with a feature cue. A computational perceptual decision-making model largely provided converging evidence that contributions from FBA were restricted to facilitating the speed with which the relevant item was identified. Our results suggest that both selection mechanisms can be used in isolation to resolve a perceptually challenging target in a sparse display, but with little additive perceptual benefit when cued simultaneously. We conclude that there is at least some higher order interdependence between space-based and feature-based selection during decision making under specific conditions.

Optimal deployment of attentional gain during fine discriminations.

Most models assume that top-down attention enhances the gain of sensory neurons tuned to behaviorally relevant stimuli (on-target gain). However, theoretical work suggests that when targets and distracters are highly similar, attention should enhance the gain of neurons that are tuned away from the target, because these neurons better discriminate neighboring features (off-target gain). While it is established that off-target neurons support difficult fine discriminations, it is unclear if top-down attentional gain can be optimally applied to informative off-target sensory neurons or if gain is always applied to on-target neurons, regardless of task demands. To test the optimality of attentional gain in human visual cortex, we used functional magnetic resonance imaging and an encoding model to estimate the response profile across a set of hypothetical orientation-selective channels during a difficult discrimination task. The results suggest that top-down attention can adaptively modulate off-target neural populations, but only when the discriminanda are precisely specified in advance. Furthermore, logistic regression revealed that activation levels in off-target orientation channels predicted behavioral accuracy on a trial-by-trial basis. Overall, these data suggest that attention does not only increase the gain of sensory-evoked responses, but may bias population response profiles in an optimal manner that respects both the tuning properties of sensory neurons and the physical characteristics of the stimulus array.

Automatic object-based spatial selection depends on the distribution of sustained attention.

Several space-based and object-based attention studies suggest these selection mechanisms may be voluntarily deployed, depending on task parameters and the attentional scope of the observer. Here, we sought to elucidate factors related to involuntary deployment of object-mediated space-based attention through two experiments. Experiment 1 used a modified flanker task where a target and nearby distractor were presented within the same or different object frames, such that an object-based attentional spread should be detrimental to performance. Results showed the presence of a flanker effect with no significant difference in magnitude between grouping conditions, indicating participants may have uniformly used a diffused attentional spotlight regardless of object segmentation. In a second experiment, we manipulated the extent of the observer's sustained attentional scope via an inducer task to determine whether object-based selection depends on the initial spotlight size. The results revealed object-based effects solely when attention narrowly encompassed the target, but not when it was widened to include the distracting flanker. This suggests the deployment of object-based attention may occur when spatial attention is initially focused narrowly. Because selecting the whole object frame directly interfered with task goals, we conclude that object-based attention may not always fully conform to relevant task goals or operate in a goal-oriented manner. We discuss these results in the context of existing literature while proposing a reconciliation of previously inconsistent findings of object-based selection.

Phasic pupillary responses modulate object-based attentional prioritization.

Visual attention studies have demonstrated that the shape of space-based selection can be governed by salient object contours: when a portion of an enclosed space is cued, the selected region extends to the full enclosure. Although this form of object-based attention (OBA) is well established, one continuing investigation focuses on whether this selection is obligatory or under voluntary control. We attempt to dissociate between these alternatives by interrogating the locus coeruleus-norepinephrine (LC-NE) system - known to fluctuate with top-down attention - during a classic two-rectangle paradigm in a sample of healthy human participants (N = 36). An endogenous spatial pre-cue directed voluntary space-based attention (SBA) to one end of a rectangular frame. We manipulated the reliability of the cue, such that targets appearing at an uncued location within the frame occurred at low or moderate frequencies. Phasic pupillary responses time-locked to the cue display served to noninvasively measure LC-NE activity, reflecting top-down processing of the spatial cue. If OBA is controlled analogously to SBA, then object selection should emerge only when it is behaviorally expedient and when LC-NE activity reflects a high degree of top-down attention to the cue display. Our results bore this out. Thus, we conclude that OBA was voluntarily controlled, and furthermore show that phasic norepinephrine may modulate attentional strategy.

Adaptive allocation of attentional gain.

Humans are adept at distinguishing between stimuli that are very similar, an ability that is particularly crucial when the outcome is of serious consequence (e.g., for a surgeon or air-traffic controller). Traditionally, selective attention was thought to facilitate perception by increasing the gain of sensory neurons tuned to the defining features of a behaviorally relevant object (e.g., color, orientation, etc.). In contrast, recent mathematical models counterintuitively suggest that, in many cases, attentional gain should be applied to neurons that are tuned away from relevant features, especially when discriminating highly similar stimuli. Here we used psychophysical methods to critically evaluate these "ideal observer" models. The data demonstrate that attention enhances the gain of the most informative sensory neurons, even when these neurons are tuned away from the behaviorally relevant target feature. Moreover, the degree to which an individual adopted optimal attentional gain settings by the end of testing predicted success rates on a difficult visual discrimination task, as well as the amount of task improvement that occurred across repeated testing sessions (learning). Contrary to most traditional accounts, these observations suggest that the primary function of attentional gain is not to enhance the representation of target features per se, but instead to optimize performance on the current perceptual task. Additionally, individual differences in gain suggest that the operating characteristics of low-level attentional phenomena are not stable trait-like attributes and that variability in how attention is deployed may play an important role in determining perceptual abilities.

Basing perceptual decisions on the most informative sensory neurons.

Single unit recording studies show that perceptual decisions are often based on the output of sensory neurons that are maximally responsive (or "tuned") to relevant stimulus features. However, when performing a difficult discrimination between two highly similar stimuli, perceptual decisions should instead be based on the activity of neurons tuned away from the relevant feature (off-channel neurons) as these neurons undergo a larger firing rate change and are thus more informative. To test this hypothesis, we measured feature-selective responses in human primary visual cortex (V1) using functional magnetic resonance imaging and show that the degree of off-channel activation predicts performance on a difficult visual discrimination task. Moreover, this predictive relationship between off-channel activation and perceptual acuity is not simply the result of extensive practice with a specific stimulus feature (as in studies of perceptual learning). Instead, relying on the output of the most informative sensory neurons may represent a general, and optimal, strategy for efficiently computing perceptual decisions.

Estimating the influence of attention on population codes in human visual cortex using voxel-based tuning functions.

In order to form stable perceptual representations, populations of sensory neurons must pool their output to overcome physiological noise; selective attention is then required to ensure that behaviorally relevant stimuli dominate these 'population codes' to gain access to awareness. However, the role that attention plays in shaping population response profiles has received little direct investigation, in part because most traditional neurophysiological methods cannot simultaneously assess changes in activity across large populations of sensory neurons. Based on single-unit recording studies, current theories hold that attending to a relevant feature sharpens the population response profile and improves the signal-to-noise ratio of the resulting perceptual representation. Here, we test this hypothesis using fMRI and an analysis approach that is able to estimate the influence of feature-based attentional modulations on population response profiles. We first derive orientation tuning functions for single voxels in human primary visual cortex, and then use these tuning functions to sort voxels according to their orientation preference. We then show that selective attention systematically biases population response profiles so that behaviorally relevant stimuli are represented in the visual system at the expense of behaviorally irrelevant stimuli. Collectively, the present results (1) provide a new approach for precisely characterizing feature-selective responses in human sensory cortices and (2) reveal how behavioral goals can shape population response profiles to support the formation of coherent perceptual representations.

Experience-dependent changes in the topography of visual crowding.

The present work examined discrimination accuracy for targets that were presented either alone in the visual field (clean displays) or embedded within a dense array of letter distractors (crowded displays). The strength of visual crowding varied strongly across the four quadrants of the visual field. Furthermore, this spatial bias in crowding was strongly influenced by the observers' prior experience with specific distractor stimuli. Observers who were monolingual readers of English experienced amplified crowding in the upper-left quadrant, while subjects with primary reading skills in Korean, Chinese, or Japanese tended towards worse target discrimination in the lower visual field. This interaction with language experience was eliminated when non-alphanumeric stimuli were employed as distractors, suggesting that prior reading experience induced a stimulus-specific change in the topography of visual crowding from English letters.

Object-based biased competition during covert spatial orienting.

Biased-competition models assert that spatial attention facilitates visual perception by biasing competitive interactions in favor of relevant input. In line with this view, past work has shown that the benefits of covert spatial attention are greatest when targets must compete with interfering stimuli. Here we propose a boundary condition for the resolution of interference via exogenous attention: Attention resolves visual interference between targets and distractors, but only when they can be individuated into distinct representations. Thus, we propose that biased competition may be object-based. We replicated previous observations of larger attention effects when targets were flanked by irrelevant distractors (interference-present displays) than when targets were presented alone (interference-absent displays). Critically, we then showed that this amplification of cueing effects in the presence of interference is eliminated when strong crowding hampers individuation of the targets and distractors. Likewise, when targets were embedded within a noise mask that did not evoke the percept of an individuated distractor, the attention effects were equivalent across noise and lone-target displays. Thus, we conclude that exogenous spatial attention resolves interference in an object-based fashion that depends on the perception of individuated targets and distractors.

Perceptual expertise enhances the resolution but not the number of representations in working memory.

Despite its central role in cognition, capacity in visual working memory is restricted to about three or four items. Curby and Gauthier (2007) examined whether perceptual expertise can help to overcome this limit by enabling more efficient coding of visual information. In line with this, they observed higher capacity estimates for upright than for inverted faces, suggesting that perceptual expertise enhances visual working memory. In the present work, we examined whether the improved capacity estimates for upright faces indicates an increased number of "slots" in working memory, or improved resolution within the existing slots. Our results suggest that perceptual expertise enhances the resolution but not the number of representations that can be held in working memory. These results clarify the effects of perceptual expertise in working memory and support recent suggestions that number and resolution represent distinct facets of working memory ability.

Spatial attention, preview, and popout: which factors influence critical spacing in crowded displays?

Crowding refers to the phenomenon in which nearby distractors impede target processing. This effect is reduced as target-distractor distance increases, and it is eliminated entirely at a distance that is labeled the critical spacing point. Attention, distractor preview, and popout are each known to facilitate processing in crowded displays. Eight experiments examined whether this is accomplished via a reduction in critical spacing. Attention was manipulated via spatial cueing, whereby a peripheral cue elicited a stimulus-driven shift of attention. Distractor preview was examined by manipulating whether the crowding distractors were presented prior to or simultaneous with the target. Popout was examined by manipulating whether there was a salient color difference between the target and distractors. As demonstrated in previous studies, we found robust benefits of spatial cueing, preview, and popout in crowded displays. However, although spatial cueing led to robust improvements in target discrimination, there was no reduction in critical spacing for attended stimuli. By contrast, both preview and popout caused large reductions in critical spacing. These disparate results indicate that attention improves target discrimination in crowded displays in a qualitatively different manner than do the other factors.

A diagnostic approach to increase reusable dinnerware selection in a cafeteria.

The current project tested a diagnostic approach to selecting interventions to increase patron selection of reusable dinnerware in a cafeteria. An assessment survey, completed by a sample of 43 patrons, suggested that the primary causes of wasteful behavior were (a) environmental arrangement of dinnerware options and (b) competing motivational variables. A functional relation between environmental arrangement and reusable product selection was demonstrated in a reversal design. However, the largest effect occurred as function of a multicomponent intervention involving environmental arrangement, employee involvement, and personal spoken prompts with motivational signs. The results support the use of informant assessments when designing community interventions.

Functions of the human frontoparietal attention network: Evidence from neuroimaging.

Human frontoparietal cortex has long been implicated as a source of attentional control. However, the mechanistic underpinnings of these control functions have remained elusive due to limitations of neuroimaging techniques that rely on anatomical landmarks to localize patterns of activation. The recent advent of topographic mapping via functional magnetic resonance imaging (fMRI) has allowed the reliable parcellation of the network into 18 independent subregions in individual subjects, thereby offering unprecedented opportunities to address a wide range of empirical questions as to how mechanisms of control operate. Here, we review the human neuroimaging literature that has begun to explore space-based, feature-based, object-based and category-based attentional control within the context of topographically defined frontoparietal cortex.

Statistical learning induces discrete shifts in the allocation of working memory resources.

Observers can voluntarily select which items are encoded into working memory, and the efficiency of this process strongly predicts memory capacity. Nevertheless, the present work suggests that voluntary intentions do not exclusively determine what is encoded into this online workspace. Observers indicated whether any items from a briefly stored sample display had changed. Unbeknown to observers, these changes were most likely to occur in a specific quadrant of the display (the dominant quadrant). Across 84 subjects and 5 groups of observers, change detection accuracy was significantly higher for items in the dominant quadrant, suggesting that memory encoding was biased towards the dominant quadrant. Only 9 of the 84 subjects were able to correctly specify the dominant quadrant when asked whether any location was more likely to contain the changed item, but more sensitive forced-choice procedures did reveal above-chance discrimination of the dominant quadrant. Nevertheless, because forced choice performance was unrelated to the size of the bias and no observer reported a biased encoding strategy, the bias was unlikely to depend on voluntary encoding strategies. The encoding bias was not due to a reduction in the response threshold for indicating changes in the dominant quadrant (Experiment 2). Finally, separate measures of the number and resolution of the representations in memory suggested that encoding was biased in a discrete slot-based fashion (Experiment 3). That is, although items in the dominant quadrant were more likely to be encoded into memory, mnemonic resolution for the favored items was not affected.

Online response-selection and the attentional blink: Multiple-processing channels.

Robust interference often arises when multiple targets (T1 and T2) are discriminated in rapid succession (the attentional blink or AB). The AB has been observed for a wide range of stimuli, and is often thought to reflect a central capacity limitation in working memory consolidation, attentional engagement, and/or online response selection. However, recent evidence challenges the existence of unitary bottleneck during postperceptual processing. Awh et al. (2004) found no AB interference when a digit target preceded a face target, presumably because these stimuli could be processed by means of separable processing channels. Using a modified AB procedure, recent studies have also demonstrated that speeded response selection of T1 leads to an AB effect for T2 identification, supporting the conclusion that response selection induces the same processing limitations that typically gives rise to an AB. The present research tests this hypothesis by examining the effects of response selection on the identification of faces. Although we replicated previous demonstrations that online response selection of a digit disrupts the identification of T2 letters, we found no interference in the identification of T2 faces. However, robust AB interference was once again observed when a speeded response to a T1 face was required, confirming that faces are not simply immune to central interference. These results dispute the existence of a unitary postperceptual capacity limitation that gives rise to the AB.