Confidence ratings do not distinguish imagination from reality.

Perceptual reality monitoring refers to the ability to distinguish internally triggered imagination from externally triggered reality. Such monitoring can take place at perceptual or cognitive levels-for example, in lucid dreaming, perceptual experience feels real but is accompanied by a cognitive insight that it is not real. We recently developed a paradigm to reveal perceptual reality monitoring errors during wakefulness in the general population, showing that imagined signals can be erroneously attributed to perception during a perceptual detection task. In the current study, we set out to investigate whether people have insight into perceptual reality monitoring errors by additionally measuring perceptual confidence. We used hierarchical Bayesian modeling of confidence criteria to characterize metacognitive insight into the effects of imagery on detection. Over two experiments, we found that confidence criteria moved in tandem with the decision criterion shift, indicating a failure of reality monitoring not only at a perceptual but also at a metacognitive level. These results further show that such failures have a perceptual rather than a decisional origin. Interestingly, offline queries at the end of the experiment revealed global, task-level insight, which was uncorrelated with local, trial-level insight as measured with confidence ratings. Taken together, our results demonstrate that confidence ratings do not distinguish imagination from reality during perceptual detection. Future research should further explore the different cognitive dimensions of insight into reality judgments and how they are related.

Dissociating the Neural Correlates of Subjective Visibility from Those of Decision Confidence.

A key goal of consciousness science is identifying neural signatures of being aware versus unaware of simple stimuli. This is often investigated in the context of near-threshold detection, with reports of stimulus awareness being linked to heightened activation in a frontoparietal network. However, because of reports of stimulus presence typically being associated with higher confidence than reports of stimulus absence, these results could be explained by frontoparietal regions encoding stimulus visibility, decision confidence, or both. In an exploratory analysis, we leverage fMRI data from 35 human participants (20 females) to disentangle these possibilities. We first show that, whereas stimulus identity was best decoded from the visual cortex, stimulus visibility (presence vs absence) was best decoded from prefrontal regions. To control for effects of confidence, we then selectively sampled trials before decoding to equalize confidence distributions between absence and presence responses. This analysis revealed striking differences in the neural correlates of subjective visibility in PFC ROIs, depending on whether or not differences in confidence were controlled for. We interpret our findings as highlighting the importance of controlling for metacognitive aspects of the decision process in the search for neural correlates of visual awareness.SIGNIFICANCE STATEMENT While much has been learned over the past two decades about the neural basis of visual awareness, the role of the PFC remains a topic of debate. By applying decoding analyses to functional brain imaging data, we show that prefrontal representations of subjective visibility are contaminated by neural correlates of decision confidence. We propose a new analysis method to control for these metacognitive aspects of awareness reports, and use it to reveal confidence-independent correlates of perceptual judgments in a subset of prefrontal areas.

Imagery adds stimulus-specific sensory evidence to perceptual detection.

Internally generated imagery and externally triggered perception rely on overlapping sensory processes. This overlap poses a challenge for perceptual reality monitoring: determining whether sensory signals reflect reality or imagination. In this study, we used psychophysics to investigate how imagery and perception interact to determine visual experience. Participants were instructed to detect oriented gratings that gradually appeared in noise while simultaneously either imagining the same grating, a grating perpendicular to the to-be-detected grating, or nothing. We found that, compared to both incongruent imagery and no imagery, congruent imagery caused a leftward shift of the psychometric function relating stimulus contrast to perceptual threshold. We discuss how this effect can best be explained by a model in which imagery adds sensory signal to the perceptual input, thereby increasing the visibility of perceived stimuli. These results suggest that, in contrast to changes in sensory signals caused by self-generated movement, the brain does not discount the influence of self-generated sensory signals on perception.

Eye movements explain decodability during perception and cued attention in MEG.

Eye movements are an integral part of human perception, but can induce artifacts in many magneto-encephalography (MEG) and electroencephalography (EEG) studies. For this reason, investigators try to minimize eye movements and remove these artifacts from their data using different techniques. When these artifacts are not purely random, but consistent regarding certain stimuli or conditions, the possibility arises that eye movements are actually inducing effects in the MEG signal. It remains unclear how much of an influence eye movements can have on observed effects in MEG, since most MEG studies lack a control analysis to verify whether an effect found in the MEG signal is induced by eye movements. Here, we find that we can decode stimulus location from eye movements in two different stages of a working memory match-to-sample task that encompass different areas of research typically done with MEG. This means that the observed MEG effect might be (partly) due to eye movements instead of any true neural correlate. We suggest how to check for eye movement effects in the data and make suggestions on how to minimize eye movement artifacts from occurring in the first place.

Vividness of Visual Imagery Depends on the Neural Overlap with Perception in Visual Areas.

Research into the neural correlates of individual differences in imagery vividness point to an important role of the early visual cortex. However, there is also great fluctuation of vividness within individuals, such that only looking at differences between people necessarily obscures the picture. In this study, we show that variation in moment-to-moment experienced vividness of visual imagery, within human subjects, depends on the activity of a large network of brain areas, including frontal, parietal, and visual areas. Furthermore, using a novel multivariate analysis technique, we show that the neural overlap between imagery and perception in the entire visual system correlates with experienced imagery vividness. This shows that the neural basis of imagery vividness is much more complicated than studies of individual differences seemed to suggest. SIGNIFICANCE STATEMENT: Visual imagery is the ability to visualize objects that are not in our direct line of sight: something that is important for memory, spatial reasoning, and many other tasks. It is known that the better people are at visual imagery, the better they can perform these tasks. However, the neural correlates of moment-to-moment variation in visual imagery remain unclear. In this study, we show that the more the neural response during imagery is similar to the neural response during perception, the more vivid or perception-like the imagery experience is.

Uncovering the Role of the Early Visual Cortex in Visual Mental Imagery.

The question of whether the early visual cortex (EVC) is involved in visual mental imagery remains a topic of debate. In this paper, I propose that the inconsistency in findings can be explained by the unique challenges associated with investigating EVC activity during imagery. During perception, the EVC processes low-level features, which means that activity is highly sensitive to variation in visual details. If the EVC has the same role during visual mental imagery, any change in the visual details of the mental image would lead to corresponding changes in EVC activity. Within this context, the question should not be whether the EVC is 'active' during imagery but how its activity relates to specific imagery properties. Studies using methods that are sensitive to variation in low-level features reveal that imagery can recruit the EVC in similar ways as perception. However, not all mental images contain a high level of visual details. Therefore, I end by considering a more nuanced view, which states that imagery can recruit the EVC, but that does not mean that it always does so.

Identifying content-invariant neural signatures of perceptual vividness.

Some conscious experiences are more vivid than others. Although perceptual vividness is a key component of human consciousness, how variation in this magnitude property is registered by the human brain is unknown. A striking feature of neural codes for magnitude in other psychological domains, such as number or reward, is that the magnitude property is represented independently of its sensory features. To test whether perceptual vividness also covaries with neural codes that are invariant to sensory content, we reanalyzed existing magnetoencephalography and functional MRI data from two distinct studies which quantified perceptual vividness via subjective ratings of awareness and visibility. Using representational similarity and decoding analyses, we find evidence for content-invariant neural signatures of perceptual vividness distributed across visual, parietal, and frontal cortices. Our findings indicate that the neural correlates of subjective vividness may share similar properties to magnitude codes in other cognitive domains.

Subjective signal strength distinguishes reality from imagination.

Humans are voracious imaginers, with internal simulations supporting memory, planning and decision-making. Because the neural mechanisms supporting imagery overlap with those supporting perception, a foundational question is how reality and imagination are kept apart. One possibility is that the intention to imagine is used to identify and discount self-generated signals during imagery. Alternatively, because internally generated signals are generally weaker, sensory strength is used to index reality. Traditional psychology experiments struggle to investigate this issue as subjects can rapidly learn that real stimuli are in play. Here, we combined one-trial-per-participant psychophysics with computational modelling and neuroimaging to show that imagined and perceived signals are in fact intermixed, with judgments of reality being determined by whether this intermixed signal is strong enough to cross a reality threshold. A consequence of this account is that when virtual or imagined signals are strong enough, they become subjectively indistinguishable from reality.

Perceptual reality monitoring: Neural mechanisms dissociating imagination from reality.

There is increasing evidence that imagination relies on similar neural mechanisms as externally triggered perception. This overlap presents a challenge for perceptual reality monitoring: deciding what is real and what is imagined. Here, we explore how perceptual reality monitoring might be implemented in the brain. We first describe sensory and cognitive factors that could dissociate imagery and perception and conclude that no single factor unambiguously signals whether an experience is internally or externally generated. We suggest that reality monitoring is implemented by higher-level cortical circuits that evaluate first-order sensory and cognitive factors to determine the source of sensory signals. According to this interpretation, perceptual reality monitoring shares core computations with metacognition. This multi-level architecture might explain several types of source confusion as well as dissociations between simply knowing whether something is real and actually experiencing it as real. We discuss avenues for future research to further our understanding of perceptual reality monitoring, an endeavour that has important implications for our understanding of clinical symptoms as well as general cognitive function.

Mistaking imagination for reality: Congruent mental imagery leads to more liberal perceptual detection.

Visual experiences can be triggered externally, by signals coming from the outside world during perception; or internally, by signals from memory during mental imagery. Imagery and perception activate similar neural codes in sensory areas, suggesting that they might sometimes be confused. In the current study, we investigated whether imagery influences perception by instructing participants to imagine gratings while externally detecting these same gratings at threshold. In a series of three experiments, we showed that imagery led to a more liberal criterion for reporting stimulus presence, and that this effect was both independent of expectation and stimulus-specific. Furthermore, participants with more vivid imagery were generally more likely to report the presence of external stimuli, independent of condition. The results can be explained as either a low-level sensory or a high-level decision-making effect. We discuss that the most likely explanation is that during imagery, internally generated sensory signals are sometimes confused for perception and suggest how the underlying mechanisms can be further characterized in future research. Our findings show that imagery and perception interact and emphasize that internally and externally generated signals are combined in complex ways to determine conscious perception.

No Evidence for Neural Overlap between Unconsciously Processed and Imagined Stimuli.

Visual representations can be generated via feedforward or feedback processes. The extent to which these processes result in overlapping representations remains unclear. Previous work has shown that imagined stimuli elicit similar representations as perceived stimuli throughout the visual cortex. However, while representations during imagery are indeed only caused by feedback processing, neural processing during perception is an interplay of both feedforward and feedback processing. This means that any representational overlap could be because of overlap in feedback processes. In the current study, we aimed to investigate this issue by characterizing the overlap between feedforward- and feedback-initiated category representations during imagined stimuli, conscious perception, and unconscious processing using fMRI in humans of either sex. While all three conditions elicited stimulus representations in left lateral occipital cortex (LOC), significant similarities were observed only between imagery and conscious perception in this area. Furthermore, connectivity analyses revealed stronger connectivity between frontal areas and left LOC during conscious perception and in imagery compared with unconscious processing. Together, these findings can be explained by the idea that long-range feedback modifies visual representations, thereby reducing representational overlap between purely feedforward- and feedback-initiated stimulus representations measured by fMRI. Neural representations influenced by feedback, either stimulus driven (perception) or purely internally driven (imagery), are, however, relatively similar.

Neural dynamics of perceptual inference and its reversal during imagery.

After the presentation of a visual stimulus, neural processing cascades from low-level sensory areas to increasingly abstract representations in higher-level areas. It is often hypothesised that a reversal in neural processing underlies the generation of mental images as abstract representations are used to construct sensory representations in the absence of sensory input. According to predictive processing theories, such reversed processing also plays a central role in later stages of perception. Direct experimental evidence of reversals in neural information flow has been missing. Here, we used a combination of machine learning and magnetoencephalography to characterise neural dynamics in humans. We provide direct evidence for a reversal of the perceptual feed-forward cascade during imagery and show that, during perception, such reversals alternate with feed-forward processing in an 11 Hz oscillatory pattern. Together, these results show how common feedback processes support both veridical perception and mental imagery.

Shared Neural Mechanisms of Visual Perception and Imagery.

For decades, the extent to which visual imagery relies on the same neural mechanisms as visual perception has been a topic of debate. Here, we review recent neuroimaging studies comparing these two forms of visual experience. Their results suggest that there is a large overlap in neural processing during perception and imagery: neural representations of imagined and perceived stimuli are similar in the visual, parietal, and frontal cortex. Furthermore, perception and imagery seem to rely on similar top-down connectivity. The most prominent difference is the absence of bottom-up processing during imagery. These findings fit well with the idea that imagery and perception rely on similar emulation or prediction processes.

Differential temporal dynamics during visual imagery and perception.

Visual perception and imagery rely on similar representations in the visual cortex. During perception, visual activity is characterized by distinct processing stages, but the temporal dynamics underlying imagery remain unclear. Here, we investigated the dynamics of visual imagery in human participants using magnetoencephalography. Firstly, we show that, compared to perception, imagery decoding becomes significant later and representations at the start of imagery already overlap with later time points. This suggests that during imagery, the entire visual representation is activated at once or that there are large differences in the timing of imagery between trials. Secondly, we found consistent overlap between imagery and perceptual processing around 160 ms and from 300 ms after stimulus onset. This indicates that the N170 gets reactivated during imagery and that imagery does not rely on early perceptual representations. Together, these results provide important insights for our understanding of the neural mechanisms of visual imagery.

Cognitive Neuroscience and Causal Inference: Implications for Psychiatry.

In this paper, we investigate to what extent it is justified to draw conclusions about causal relations between brain states and mental states from cognitive neuroscience studies. We first explain the views of two prominent proponents of the interventionist account of causation: Woodward and Baumgartner. We then discuss the implications of their views in the context of traditional cognitive neuroscience studies in which the effect of changes in mental state on changes in brain states is investigated. After this, we turn to brain stimulation studies in which brain states are manipulated to investigate the effects on mental states. We argue that, depending on whether one sides with Woodward or Baumgartner, it is possible to draw causal conclusions from both types of studies (Woodward) or from brain stimulation studies only (Baumgartner). We show what happens to these conclusions if we adopt different views of the relation between mental states and brain states. Finally, we discuss the implications of our findings for psychiatry and the treatment of psychiatric disorders.

The spatiotemporal dynamics of binocular rivalry: evidence for increased top-down flow prior to a perceptual switch.

According to most theories, perceptual switching during binocular rivalry is caused by competition between the neural representations of the two input images. It remains unclear whether competition is resolved already at the early stages of visual processing and that information about the dominant percept is then fed forward to more high-level areas or whether competition is first resolved in high-level areas and then fed back to lower levels. This study aimed to dissociate between these theories by investigating the direction of information flow prior to a perceptual switch, using Granger causality on classifier output originating from occipital, temporal, parietal and frontal regions of interest. The results point toward increased top-down information flow between temporal and occipital areas before a switch in dominance. These findings do not support a low-level account of binocular rivalry but are in line with high-level and hybrid explanations.