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.

Prior Expectations of Motion Direction Modulate Early Sensory Processing.

Perception is a process of inference, integrating sensory inputs with prior expectations. However, little is known regarding the temporal dynamics of this integration. It has been proposed that expectation plays a role early in the perceptual process, biasing sensory processing. Alternatively, others suggest that expectations are integrated only at later, postperceptual decision-making stages. The current study aimed to dissociate between these hypotheses. We exposed human participants (male and female) to auditory cues predicting the likely direction of upcoming moving dot patterns, while recording neural activity using magnetoencephalography (MEG). Participants' reports of the moving dot directions were biased toward the direction predicted by the cues. To investigate when expectations affected sensory representations, we used inverted encoding models to decode the direction represented in early sensory signals. Strikingly, the cues modulated the direction represented in the MEG signal as early as 150 ms after visual stimulus onset. While this may not reflect a modulation of the initial feedforward sweep, it does reveal a modulation of early sensory representations. Exploratory analyses showed that the neural modulation was related to perceptual expectation effects: participants with a stronger perceptual bias toward the predicted direction also revealed a stronger reflection of the predicted direction in the MEG signal. For participants with this perceptual bias, a correlation between decoded and perceived direction already emerged before visual stimulus onset, suggesting that the prestimulus state of the visual cortex influences sensory processing. Together, these results suggest that expectations play an integral role in the neural computations underlying perception.SIGNIFICANCE STATEMENT Perception can be thought of as an inferential process in which our brains integrate sensory inputs with prior expectations to make sense of the world. This study investigated whether this integration occurs early or late in the process of perception. We exposed human participants to auditory cues that predicted the likely direction of visual moving dots, while recording neural activity with millisecond resolution using magnetoencephalography. Participants' perceptual reports of the direction of the moving dots were biased toward the predicted direction. Additionally, the predicted direction modulated the neural representation of the moving dots just 150 ms after they appeared. This suggests that prior expectations affected sensory processing at early stages, playing an integral role in the perceptual process.

Prior expectations evoke stimulus templates in the primary visual cortex.

Sensory processing is strongly influenced by prior expectations. Valid expectations have been shown to lead to improvements in perception as well as in the quality of sensory representations in primary visual cortex. However, very little is known about the neural correlates of the expectations themselves. Previous studies have demonstrated increased activity in sensory cortex following the omission of an expected stimulus, yet it is unclear whether this increased activity constitutes a general surprise signal or rather has representational content. One intriguing possibility is that top-down expectation leads to the formation of a template of the expected stimulus in visual cortex, which can then be compared with subsequent bottom-up input. To test this hypothesis, we used fMRI to noninvasively measure neural activity patterns in early visual cortex of human participants during expected but omitted visual stimuli. Our results show that prior expectation of a specific visual stimulus evokes a feature-specific pattern of activity in the primary visual cortex (V1) similar to that evoked by the corresponding actual stimulus. These results are in line with the notion that prior expectation triggers the formation of specific stimulus templates to efficiently process expected sensory inputs.

Shared representations for working memory and mental imagery in early visual cortex.

Early visual areas contain specific information about visual items maintained in working memory, suggesting a role for early visual cortex in more complex cognitive functions [1-4]. It is an open question, however, whether these areas also underlie the ability to internally generate images de novo (i.e., mental imagery). Research on mental imagery has to this point focused mostly on whether mental images activate early sensory areas, with mixed results [5-7]. Recent studies suggest that multivariate pattern analysis of neural activity patterns in visual regions can reveal content-specific representations during cognitive processes, even though overall activation levels are low [1-4]. Here, we used this approach [8, 9] to study item-specific activity patterns in early visual areas (V1-V3) when these items are internally generated. We could reliably decode stimulus identity from neural activity patterns in early visual cortex during both working memory and mental imagery. Crucially, these activity patterns resembled those evoked by bottom-up visual stimulation, suggesting that mental images are indeed "perception-like" in nature. These findings suggest that the visual cortex serves as a dynamic "blackboard" [10, 11] that is used during both bottom-up stimulus processing and top-down internal generation of mental content.