Non-visual spatial strategies are effective for maintaining precise information in visual working memory.

Visual working memory content is commonly thought to be composed of a precise visual representation of stimulus information (e.g., color, shape). Nevertheless, previous research has shown that individuals represent this visual information in different formats, historically dichotomized into "verbal" and "visual" formats. With growing popular knowledge of aphantasia, or the absence of sensory mental imagery, recent studies have demonstrated that individuals with aphantasia perform similarly to individuals with typical imagery on visual working memory tasks. This suggest that the use of non-visual strategies may be sufficient to perform visual working memory tasks, which were previously thought to be strictly visual. To investigate the effects of different strategies on performance in a visual working memory task, we recruited individuals across the visual imagery spectrum and tested their ability to identify relatively small (3°), medium (6°), or large (10°) changes in the degree of orientation of gratings held in working memory. Subsequently, participants indicated the extent to which they used five different strategies: visual, spatial, verbal, semantic, and sensorimotor. Results revealed that individuals with aphantasia and typical imagery performed similarly to each other across all task difficulty levels. Individuals with typical imagery dominantly used visuospatial strategies, but surprisingly, individuals with aphantasia overwhelmingly preferred the use of non-visual spatial and sensorimotor strategies over verbal strategies. These results suggest that non-visual spatial and sensorimotor strategies can be adopted in visual working memory tasks and these strategies are equally effective as visuospatial strategies. This calls for a rethinking of the "visual" versus "verbal" dichotomy, and provides evidence for the use of other non-visual mental representations in working memory tasks.

A novel model of divergent predictive perception.

Predictive processing theories state that our subjective experience of reality is shaped by a balance of expectations based on previous knowledge about the world (i.e. priors) and confidence in sensory input from the environment. Divergent experiences (e.g. hallucinations and synaesthesia) are likely to occur when there is an imbalance between one's reliance on priors and sensory input. In a novel theoretical model, inspired by both predictive processing and psychological principles, we propose that predictable divergent experiences are associated with natural or environmentally induced prior/sensory imbalances: inappropriately strong or inflexible (i.e. maladaptive) high-level priors (beliefs) combined with low sensory confidence can result in reality discrimination issues, a characteristic of psychosis; maladaptive low-level priors (sensory expectations) combined with high sensory confidence can result in atypical sensory sensitivities and persistent divergent percepts, a characteristic of synaesthesia. Crucially, we propose that whether different divergent experiences manifest with dominantly sensory (e.g. hallucinations) or nonsensory characteristics (e.g. delusions) depends on mental imagery ability, which is a spectrum from aphantasia (absent or weak imagery) to hyperphantasia (extremely vivid imagery). We theorize that imagery is critically involved in shaping the sensory richness of divergent perceptual experience. In sum, to predict a range of divergent perceptual experiences in both clinical and general populations, three factors must be accounted for: a maladaptive use of priors, individual level of confidence in sensory input, and mental imagery ability. These ideas can be expressed formally using nonparametric regression modeling. We provide evidence for our theory from previous work and deliver predictions for future research.

The Ganzflicker experience: High probability of seeing vivid and complex pseudo-hallucinations with imagery but not aphantasia.

There are considerable individual differences in visual mental imagery ability across the general population, including a "blind mind's eye", or aphantasia. Recent studies have shown that imagery is linked to differences in perception in the healthy population, and clinical work has found a connection between imagery and hallucinatory experiences in neurological disorders. However, whether imagery ability is associated with anomalous perception-including hallucinations-in the general population remains unclear. In the current study, we explored the relationship between imagery ability and the anomalous perception of pseudo-hallucinations (PH) using rhythmic flicker stimulation ("Ganzflicker"). Specifically, we investigated whether the ability to generate voluntary imagery is associated with susceptibility to flicker-induced PH. We additionally explored individual differences in observed features of PH. We recruited a sample of people with aphantasia (aphants) and imagery (imagers) to view a constant red-and-black flicker for approximately 10 min. We found that imagers were more susceptible to PH, and saw more complex and vivid PH, compared to aphants. This study provides the first evidence that the ability to generate visual imagery increases the likelihood of experiencing complex and vivid anomalous percepts.

Anomalous visual experience is linked to perceptual uncertainty and visual imagery vividness.

An imbalance between top-down and bottom-up processing on perception (specifically, over-reliance on top-down processing) can lead to anomalous perception, such as illusions. One factor that may be involved in anomalous perception is visual mental imagery, which is the experience of "seeing" with the mind's eye. There are vast individual differences in self-reported imagery vividness, and more vivid imagery is linked to a more sensory-like experience. We, therefore, hypothesized that susceptibility to anomalous perception is linked to individual imagery vividness. To investigate this, we adopted a paradigm that is known to elicit the perception of faces in pure visual noise (pareidolia). In four experiments, we explored how imagery vividness contributes to this experience under different response instructions and environments. We found strong evidence that people with more vivid imagery were more likely to see faces in the noise, although removing suggestive instructions weakened this relationship. Analyses from the first two experiments led us to explore confidence as another factor in pareidolia proneness. We, therefore, modulated environment noise and added a confidence rating in a novel design. We found strong evidence that pareidolia proneness is correlated with uncertainty about real percepts. Decreasing perceptual ambiguity abolished the relationship between pareidolia proneness and both imagery vividness and confidence. The results cannot be explained by incidental face-like patterns in the noise, individual variations in response bias, perceptual sensitivity, subjective perceptual thresholds, viewing distance, testing environments, motivation, gender, or prosopagnosia. This indicates a critical role of mental imagery vividness and perceptual uncertainty in anomalous perceptual experience.

Diagnostic parts are not exclusive in the search template for real-world object categories.

Visual search can be aided by a search template: a preparatory representation of relevant target features. But which features are relevant in complex, real-world category search? Previous research suggests that this template must be flexible to account for variations in naturalistic stimulus properties such as size and occlusion, and that shapes of diagnostic parts of objects are a likely candidate. Here, in three experiments, we systematically evaluated the contribution of diagnostic object parts and whole object shape to the category-level search template. Our hypothesis was that features that better match the active search template will capture attention during search more strongly than partially-matching features. Results showed that while whole objects captured attention reliably and globally across the visual field, diagnostic parts failed to do so in all three experiments. This suggests that whole object shape is a necessary component of the category-level search template.

Task relevance modulates the cortical representation of feature conjunctions in the target template.

Little is known about the cortical regions involved in representing task-related content in preparation for visual task performance. Here we used representational similarity analysis (RSA) to investigate the BOLD response pattern similarity between task relevant and task irrelevant feature dimensions during conjunction viewing and target template maintenance prior to visual search. Subjects were cued to search for a spatial frequency (SF) or orientation of a Gabor grating and we measured BOLD signal during cue and delay periods before the onset of a search display. RSA of delay period activity revealed that widespread regions in frontal, posterior parietal, and occipitotemporal cortices showed general representational differences between task relevant and task irrelevant dimensions (e.g., orientation vs. SF). In contrast, RSA of cue period activity revealed sensory-related representational differences between cue images (regardless of task) at the occipital pole and additionally in the frontal pole. Our data show that task and sensory information are represented differently during viewing and during target template maintenance, and that task relevance modulates the representation of visual information across the cortex.

Individual differences shape the content of visual representations.

Visually perceiving a stimulus activates a pictorial representation of that item in the brain, but how pictorial is the representation of a stimulus in the absence of visual stimulation? Here I address this question with a review of the literatures on visual imagery (VI), visual working memory (VWM), and visual preparatory templates, all of which require activating visual information in the absence of sensory stimulation. These processes have historically been studied separately, but I propose that they can provide complimentary evidence for the pictorial nature of their contents. One major challenge in studying the contents of visual representations is the discrepant findings concerning the extent of overlap (both cortical and behavioral) between externally and internally sourced visual representations. I argue that these discrepancies may in large part be due to individual differences in VI vividness and precision, the specific representative abilities required to perform a task, appropriateness of visual preparatory strategies, visual cortex anatomy, and level of expertise with a particular object category. Individual differences in visual representative abilities greatly impact task performance and may influence the likelihood of experiences such as intrusive VI and hallucinations, but research still predominantly focuses on uniformities in visual experience across individuals. In this paper I review the evidence for the pictorial content of visual representations activated for VI, VWM, and preparatory templates, and highlight the importance of accounting for various individual differences in conducting research on this topic.

Privileged access to awareness for faces and objects of expertise.

Access to visual awareness for human faces is strongly influenced by spatial orientation: Under continuous flash suppression (CFS), upright faces break into awareness more quickly than inverted faces. This effect of inversion for faces is larger than for a wide range of other animate and inanimate objects. Here we asked whether this apparently specific sensitivity to upright faces reflects face-specific detection mechanisms or whether it reflects perceptual expertise more generally. We tested car experts who varied in their degree of car and face expertise and measured the time upright and inverted faces, cars, and chairs needed to overcome CFS and break into awareness. Results showed that greater car expertise was correlated with larger car inversion effects under CFS. A similar relation between better discrimination performance and larger CFS inversion effects was found for faces. CFS inversion effects are thus modulated by perceptual expertise for both faces and cars. These results demonstrate that inversion effects in conscious access are not unique to faces but similarly exist for other objects of expertise. More generally, we interpret these findings as suggesting that access to awareness and exemplar-level discrimination rely on partially shared perceptual mechanisms. (PsycINFO Database Record

Perceptual expertise improves category detection in natural scenes.

There is much debate about how detection, categorization, and within-category identification relate to one another during object recognition. Whether these tasks rely on partially shared perceptual mechanisms may be determined by testing whether training on one of these tasks facilitates performance on another. In the present study we asked whether expertise in discriminating objects improves the detection of these objects in naturalistic scenes. Self-proclaimed car experts (N = 34) performed a car discrimination task to establish their level of expertise, followed by a visual search task where they were asked to detect cars and people in hundreds of photographs of natural scenes. Results revealed that expertise in discriminating cars was strongly correlated with car detection accuracy. This effect was specific to objects of expertise, as there was no influence of car expertise on person detection. These results indicate a close link between object discrimination and object detection performance, which we interpret as reflecting partially shared perceptual mechanisms and neural representations underlying these tasks: the increased sensitivity of the visual system for objects of expertise - as a result of extensive discrimination training - may benefit both the discrimination and the detection of these objects. Alternative interpretations are also discussed.

Preparatory Activity in Posterior Temporal Cortex Causally Contributes to Object Detection in Scenes.

Theories of visual selective attention propose that top-down preparatory attention signals mediate the selection of task-relevant information in cluttered scenes. Neuroimaging and electrophysiology studies have provided correlative evidence for this hypothesis, finding increased activity in target-selective neural populations in visual cortex in the period between a search cue and target onset. In this study, we used online TMS to test whether preparatory neural activity in visual cortex is causally involved in naturalistic object detection. In two experiments, participants detected the presence of object categories (cars, people) in a diverse set of photographs of real-world scenes. TMS was applied over a region in posterior temporal cortex identified by fMRI as carrying category-specific preparatory activity patterns. Results showed that TMS applied over posterior temporal cortex before scene onset (-200 and -100 msec) impaired the detection of object categories in subsequently presented scenes, relative to vertex and early visual cortex stimulation. This effect was specific to category level detection and was related to the type of attentional template participants adopted, with the strongest effects observed in participants adopting category level templates. These results provide evidence for a causal role of preparatory attention in mediating the detection of objects in cluttered daily-life environments.

Involuntary attentional capture by task-irrelevant objects that match the search template for category detection in natural scenes.

Theories of visual search postulate that the selection of targets amongst distractors involves matching visual input to a top-down attentional template. Previous work has provided evidence that feature-based attentional templates affect visual processing globally across the visual field. In the present study, we asked whether more naturalistic, category-level attentional templates also modulate visual processing in a spatially global and obligatory way. Subjects were cued to detect people or cars in a diverse set of photographs of real-world scenes. On a subset of trials, silhouettes of people and cars appeared in search-irrelevant locations that subjects were instructed to ignore, and subjects were required to respond to the location of a subsequent dot probe. In three experiments, results showed a consistency effect on dot-probe trials: dot probes were detected faster when they appeared in the location of the cued category compared with the non-cued category, indicating attentional capture by template-matching stimuli. Experiments 1 and 2 showed that this capture was involuntary: consistency effects persisted under conditions in which attending to silhouettes of the cued category was detrimental to performance. Experiment 3 tested whether these effects could be attributed to non-attentional effects related to the processing of the category cues. Results showed a consistency effect when subjects searched for category exemplars but not when they searched for objects semantically related to the cued category. Together, these results indicate that attentional templates for familiar object categories affect visual processing across the visual field, leading to involuntary attentional capture by template-matching stimuli.

The contents of the search template for category-level search in natural scenes.

Visual search involves the matching of visual input to a "search template," an internal representation of task-relevant information. The present study investigated the contents of the search template during visual search for object categories in natural scenes, for which low-level features do not reliably distinguish targets from nontargets. Subjects were cued to detect people or cars in diverse photographs of real-world scenes. On a subset of trials, the cue was followed by task-irrelevant stimuli instead of scenes, directly followed by a dot that subjects were instructed to detect. We hypothesized that stimuli that matched the active search template would capture attention, resulting in faster detection of the dot when presented at the location of a template-matching stimulus. Results revealed that silhouettes of cars and people captured attention irrespective of their orientation (0°, 90°, or 180°). Interestingly, strong capture was observed for silhouettes of category-diagnostic object parts, such as the wheel of a car. Finally, attentional capture was also observed for silhouettes presented at locations that were irrelevant to the search task. Together, these results indicate that search for familiar object categories in real-world scenes is mediated by spatially global search templates that consist of view-invariant shape representations of category-diagnostic object parts.