Slower but more accurate mental rotation performance in aphantasia linked to differences in cognitive strategies.

Mental rotation tasks are frequently used as standard measures of mental imagery. However, aphantasia research has brought such use into question. Here, we assessed a large group of individuals who lack visual imagery (aphantasia) on two mental rotation tasks: a three-dimensional block-shape, and a human manikin rotation task. In both tasks, those with aphantasia had slower, but more accurate responses than controls. Both groups demonstrated classic linear increases in response time and error-rate as functions of angular disparity. In the three-dimensional block-shape rotation task, a within-group speed-accuracy trade-off was found in controls, whereas faster individuals in the aphantasia group were also more accurate. Control participants generally favoured using object-based mental rotation strategies, whereas those with aphantasia favoured analytic strategies. These results suggest that visual imagery is not crucial for successful performance in classical mental rotation tasks, as alternative strategies can be effectively utilised in the absence of holistic mental representations.

Revisiting the blind mind: Still no evidence for sensory visual imagery in individuals with aphantasia.

The inability to visualise was given the name aphantasia in 2015 by Zeman and colleagues. In 2018 we published research showing that fifteen individuals who self-identified as having aphantasia also demonstrated a lack of sensory visual imagery when undergoing the binocular rivalry imagery paradigm, suggesting more than just a metacognitive difference. Here we update these findings with over fifty participants with aphantasia and show that there is evidence for a lack of sensory imagery in aphantasia. How the binocular rivalry paradigm scores relate to the vividness of visual imagery questionnaire (VVIQ) and how aphantasia can be confirmed is discussed.

Multisensory subtypes of aphantasia: Mental imagery as supramodal perception in reverse.

Cognitive neuroscience research on mental imagery has largely focused on the visual imagery modality in unimodal task contexts. Recent studies have uncovered striking individual differences in visual imagery capacity, with some individuals reporting a subjective absence of conscious visual imagery ability altogether ("aphantasia"). However, naturalistic mental imagery is often multi-sensory, and preliminary findings suggest that many individuals with aphantasia also report a subjective lack of mental imagery in other sensory domains (such as auditory or olfactory imagery). In this paper, we perform a series of cluster analyses on the multi-sensory imagery questionnaire scores of two large groups of aphantasic subjects, defining latent sub-groups in this sample population. We demonstrate that aphantasia is a heterogenous phenomenon characterised by dominant sub-groups of individuals with visual aphantasia (those who report selective visual imagery absence) and multi-sensory aphantasia (those who report an inability to generate conscious mental imagery in any sensory modality). We replicate our findings in a second large sample and show that more unique aphantasia sub-types also exist, such as individuals with selectively preserved mental imagery in only one sensory modality (e.g. intact auditory imagery). We outline the implications of our findings for network theories of mental imagery, discussing how unique aphantasia aetiologies with distinct self-report patterns might reveal alterations to various levels of the sensory processing hierarchy implicated in mental imagery.

Different Mechanisms for Supporting Mental Imagery and Perceptual Representations: Modulation Versus Excitation.

Recent research suggests imagery is functionally equivalent to a weak form of visual perception. Here we report evidence across five independent experiments on adults that perception and imagery are supported by fundamentally different mechanisms: Whereas perceptual representations are largely formed via increases in excitatory activity, imagery representations are largely supported by modulating nonimagined content. We developed two behavioral techniques that allowed us to first put the visual system into a state of adaptation and then probe the additivity of perception and imagery. If imagery drives similar excitatory visual activity to perception, pairing imagery with perceptual adapters should increase the state of adaptation. Whereas pairing weak perception with adapters increased measures of adaptation, pairing imagery reversed their effects. Further experiments demonstrated that these nonadditive effects were due to imagery weakening representations of nonimagined content. Together these data provide empirical evidence that the brain uses categorically different mechanisms to represent imagery and perception.

Memories with a blind mind: Remembering the past and imagining the future with aphantasia.

Our capacity to re-experience the past and simulate the future is thought to depend heavily on visual imagery, which allows us to construct complex sensory representations in the absence of sensory stimulation. There are large individual differences in visual imagery ability, but their impact on autobiographical memory and future prospection remains poorly understood. Research in this field assumes the normative use of visual imagery as a cognitive tool to simulate the past and future, however some individuals lack the ability to visualise altogether (a condition termed "aphantasia"). Aphantasia represents a rare and naturally occurring knock-out model for examining the role of visual imagery in episodic memory recall. Here, we assessed individuals with aphantasia on an adapted form of the Autobiographical Interview, a behavioural measure of the specificity and richness of episodic details underpinning the memory of events. Aphantasic participants generated significantly fewer episodic details than controls for both past and future events. This effect was most pronounced for novel future events, driven by selective reductions in visual detail retrieval, accompanied by comparatively reduced ratings of the phenomenological richness of simulated events, and paralleled by quantitative linguistic markers of reduced perceptual language use in aphantasic participants compared to those with visual imagery. Our findings represent the first systematic evidence (using combined objective and subjective data streams) that aphantasia is associated with a diminished ability to re-experience the past and simulate the future, indicating that visual imagery is an important cognitive tool for the dynamic retrieval and recombination of episodic details during mental simulation.

Aphantasia, dysikonesia, anauralia: call for a single term for the lack of mental imagery-Commentary on Dance et al. (2021) and Hinwar and Lambert (2021).

Recently, the term 'aphantasia' has become current in scientific and public discourse to denote the absence of mental imagery. However, new terms for aphantasia or its subgroups have recently been proposed, e.g., 'dysikonesia' or 'anauralia', which complicates the literature, research communication and understanding for the general public. Before further terms emerge, we advocate the consistent use of the term 'aphantasia' as it can be used flexibly and precisely, and is already widely known in the scientific community and among the general public.

The pupillary light response as a physiological index of aphantasia, sensory and phenomenological imagery strength.

The pupillary light response is an important automatic physiological response which optimizes light reaching the retina. Recent work has shown that the pupil also adjusts in response to illusory brightness and a range of cognitive functions, however, it remains unclear what exactly drives these endogenous changes. Here, we show that the imagery pupillary light response correlates with objective measures of sensory imagery strength. Further, the trial-by-trial phenomenological vividness of visual imagery is tracked by the imagery pupillary light response. We also demonstrated that a group of individuals without visual imagery (aphantasia) do not show any significant evidence of an imagery pupillary light response, however they do show perceptual pupil light responses and pupil dilation with larger cognitive load. Our results provide evidence that the pupillary light response indexes the sensory strength of visual imagery. This work also provides the first physiological validation of aphantasia.

Visual working memory in aphantasia: Retained accuracy and capacity with a different strategy.

Visual working memory paradigms involve retaining and manipulating visual information in mind over a period of seconds. Evidence suggests that visual imagery (sensory recruitment) is a strategy used by many to retain visual information during such tasks, leading some researchers to propose that visual imagery and visual working memory may be one and the same. If visual imagery is essential to visual working memory task performance there should be large ramifications for a special population of individuals who do not experience visual imagery, aphantasia. Here we assessed visual working memory task performance in this population using a number of different lab and clinical working memory tasks. We found no differences in capacity limits for visual, general number or spatial working memory for aphantasic individuals compared to controls. Further, aphantasic individuals showed no significant differences in performance on visual components of clinical working memory tests as compared to verbal components. However, there were significant differences in the reported strategies used by aphantasic individuals across all memory tasks. Additionally, aphantasic individual's visual memory accuracy did not demonstrate a significant oblique orientation effect, which is proposed to occur due to sensory recruitment, further supporting their non-visual imagery strategy reports. Taken together these data demonstrate that aphantasic individuals are not impaired on visual working memory tasks, suggesting visual imagery and working memory are not one and the same, with imagery (and sensory recruitment) being just one of the tools that can be used to solve visual working memory tasks.

Aphantasia: The science of visual imagery extremes.

Visual imagery allows us to revisit the appearance of things in their absence and to test out virtual combinations of sensory experience. Visual imagery has been linked to many cognitive processes, such as autobiographical and visual working memory. Imagery also plays symptomatic and mechanistic roles in neurologic and mental disorders and is utilized in treatment. A large network of brain activity spanning frontal, parietal, temporal, and visual cortex is involved in generating and maintain images in mind. The ability to visualize has extreme variations, ranging from completely absent (aphantasia) to photo-like (hyperphantasia). The anatomy and functionality of visual cortex, including primary visual cortex, have been associated with individual differences in visual imagery ability, pointing to a potential correlate for both aphantasia and hyperphantasia. Preliminary evidence suggests that lifelong aphantasia is associated with prosopagnosia and reduction in autobiographical memory; hyperphantasia is associated with synesthesia. Aphantasic individuals can also be highly imaginative and are able to complete many tasks that were previously thought to rely on visual imagery, demonstrating that visualization is only one of many ways of representing things in their absence. The study of extreme imagination reminds us how easily invisible differences can escape detection.

The critical role of mental imagery in human emotion: insights from fear-based imagery and aphantasia.

One proposed function of imagery is to make thoughts more emotionally evocative through sensory simulation, which can be helpful both in planning for future events and in remembering the past, but also a hindrance when thoughts become overwhelming and maladaptive, such as in anxiety disorders. Here, we report a novel test of this theory using a special population with no visual imagery: aphantasia. After using multi-method verification of aphantasia, we show that this condition, but not the general population, is associated with a flat-line physiological response (skin conductance levels) to reading and imagining frightening stories. Importantly, we show in a second experiment that this difference in physiological responses to fear-inducing stimuli is not found when perceptually viewing fearful images. These data demonstrate that the aphantasic individuals' lack of a physiological response when imaging scenarios is likely to be driven by their inability to visualize and is not due to a general emotional or physiological dampening. This work provides evidence that a lack of visual imagery results in a dampened emotional response when reading fearful scenarios, providing evidence for the emotional amplification theory of visual imagery.

Why do imagery and perception look and feel so different?

Despite the past few decades of research providing convincing evidence of the similarities in function and neural mechanisms between imagery and perception, for most of us, the experience of the two are undeniably different, why? Here, we review and discuss the differences between imagery and perception and the possible underlying causes of these differences, from function to neural mechanisms. Specifically, we discuss the directional flow of information (top-down versus bottom-up), the differences in targeted cortical layers in primary visual cortex and possible different neural mechanisms of modulation versus excitation. For the first time in history, neuroscience is beginning to shed light on this long-held mystery of why imagery and perception look and feel so different. This article is part of the theme issue 'Offline perception: voluntary and spontaneous perceptual experiences without matching external stimulation'.

Attention driven phantom vision: measuring the sensory strength of attentional templates and their relation to visual mental imagery and aphantasia.

When we search for an object in an array or anticipate attending to a future object, we create an 'attentional template' of the object. The definitions of attentional templates and visual imagery share many similarities as well as many of the same neural characteristics. However, the phenomenology of these attentional templates and their neural similarities to visual imagery and perception are rarely, if ever discussed. Here, we investigate the relationship between these two forms of non-retinal phantom vision through the use of the binocular rivalry technique, which allows us to measure the sensory strength of attentional templates in the absence of concurrent perceptual stimuli. We find that attentional templates correlate with both feature-based attention and visual imagery. Attentional templates, like imagery, were significantly disrupted by the presence of irrelevant visual stimuli, while feature-based attention was not. We also found that a special population who lack the ability to visualize (aphantasia), showed evidence of feature-based attention when measured using the binocular rivalry paradigm, but not attentional templates. Taken together, these data suggest functional similarities between attentional templates and visual imagery, advancing the theory of visual imagery as a general simulation tool used across cognition. This article is part of the theme issue 'Offline perception: voluntary and spontaneous perceptual experiences without matching external stimulation'.

Offline perception: an introduction.

Experiences that are self-generated and independent of sensory stimulations permeate our whole life. This theme issue examines their similarities and differences, systematizes the literature from an integrative perspective, critically discusses state-of-the-art empirical findings and proposes new theoretical approaches. The aim of the theme issue is to foster interaction between the different disciplines and research directions involved and to explore the prospects of a unificatory account of offline perception in general. This article is part of the theme issue 'Offline perception: voluntary and spontaneous perceptual experiences without matching external stimulation'.

Decoding Nonconscious Thought Representations during Successful Thought Suppression.

Controlling our thoughts is central to mental well-being, and its failure is at the crux of a number of mental disorders. Paradoxically, behavioral evidence shows that thought suppression often fails. Despite the broad importance of understanding the mechanisms of thought control, little is known about the fate of neural representations of suppressed thoughts. Using fMRI, we investigated the brain areas involved in controlling visual thoughts and tracked suppressed thought representations using multivoxel pattern analysis. Participants were asked to either visualize a vegetable/fruit or suppress any visual thoughts about those objects. Surprisingly, the content (object identity) of successfully suppressed thoughts was still decodable in visual areas with algorithms trained on imagery. This suggests that visual representations of suppressed thoughts are still present despite reports that they are not. Thought generation was associated with the left hemisphere, and thought suppression was associated with right hemisphere engagement. Furthermore, general linear model analyses showed that subjective success in thought suppression was correlated with engagement of executive areas, whereas thought-suppression failure was associated with engagement of visual and memory-related areas. These results suggest that the content of suppressed thoughts exists hidden from awareness, seemingly without an individual's knowledge, providing a compelling reason why thought suppression is so ineffective. These data inform models of unconscious thought production and could be used to develop new treatment approaches to disorders involving maladaptive thoughts.

A cognitive profile of multi-sensory imagery, memory and dreaming in aphantasia.

For most people, visual imagery is an innate feature of many of our internal experiences, and appears to play a critical role in supporting core cognitive processes. Some individuals, however, lack the ability to voluntarily generate visual imagery altogether - a condition termed "aphantasia". Recent research suggests that aphantasia is a condition defined by the absence of visual imagery, rather than a lack of metacognitive awareness of internal visual imagery. Here we further illustrate a cognitive "fingerprint" of aphantasia, demonstrating that compared to control participants with imagery ability, aphantasic individuals report decreased imagery in other sensory domains, although not all report a complete lack of multi-sensory imagery. They also report less vivid and phenomenologically rich autobiographical memories and imagined future scenarios, suggesting a constructive role for visual imagery in representing episodic events. Interestingly, aphantasic individuals report fewer and qualitatively impoverished dreams compared to controls. However, spatial abilities appear unaffected, and aphantasic individuals do not appear to be considerably protected against all forms of trauma symptomatology in response to stressful life events. Collectively, these data suggest that imagery may be a normative representational tool for wider cognitive processes, highlighting the large inter-individual variability that characterises our internal mental representations.

Cortical excitability controls the strength of mental imagery.

Mental imagery provides an essential simulation tool for remembering the past and planning the future, with its strength affecting both cognition and mental health. Research suggests that neural activity spanning prefrontal, parietal, temporal, and visual areas supports the generation of mental images. Exactly how this network controls the strength of visual imagery remains unknown. Here, brain imaging and transcranial magnetic phosphene data show that lower resting activity and excitability levels in early visual cortex (V1-V3) predict stronger sensory imagery. Further, electrically decreasing visual cortex excitability using tDCS increases imagery strength, demonstrating a causative role of visual cortex excitability in controlling visual imagery. Together, these data suggest a neurophysiological mechanism of cortical excitability involved in controlling the strength of mental images.

The functional effects of voluntary and involuntary phantom color on conscious awareness.

The constructive nature of vision is perhaps most evident during hallucinations, mental imagery, synesthesia, perceptual filling-in, and many illusions in which conscious visual experience does not overtly correspond to retinal stimulation: phantom vision. However, the relationship between voluntary and involuntary phantom vision remains largely unknown. Here, we investigated 2 forms of visual phantom color, neon phantom color spreading and voluntary color mental imagery, and their effect on subsequent binocular rivalry perception. Passively viewing neon phantom color induced time sensitive, suppressive effects on subsequent binocular rivalry. These effects could be attenuated by rotating the color inducers, or like color imagery, by concurrent uniform luminance stimulation. The degree of neon color induced rivalry suppression predicted the degree of voluntary color imagery facilitation, both on subsequent rivalry perception. Further, there were functional interactions between voluntary imagery and involuntary neon phantom vision when experienced successively. Our results suggest potential sensory mechanistic commonalities between voluntary and involuntary phantom vision. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

The human imagination: the cognitive neuroscience of visual mental imagery.

Mental imagery can be advantageous, unnecessary and even clinically disruptive. With methodological constraints now overcome, research has shown that visual imagery involves a network of brain areas from the frontal cortex to sensory areas, overlapping with the default mode network, and can function much like a weak version of afferent perception. Imagery vividness and strength range from completely absent (aphantasia) to photo-like (hyperphantasia). Both the anatomy and function of the primary visual cortex are related to visual imagery. The use of imagery as a tool has been linked to many compound cognitive processes and imagery plays both symptomatic and mechanistic roles in neurological and mental disorders and treatments.