Prime-induced illusion of control: The influence of unconscious priming on self-initiated actions and the role of regression to the mean.

To what degree human cognition is influenced by subliminal stimuli is a controversial empirical question. One striking example was reported by Linser and Goschke (2007): participants overestimated how much control they had over objectively uncontrollable stimuli when masked congruent primes were presented immediately before the action. Critically, however, unawareness of the masked primes was established by post hoc data selection. In our preregistered study we sought to explore these findings while adjusting prime visibility based on individual thresholds, so that each participant underwent both visible and non-visible conditions. In experiment 1, N = 39 participants engaged in a control judgement task: following the presentation of a semantic prime, they freely selected between two keys, which triggered the appearance of a colored circle. The color of the circles, however, was independent of the key-press. Subsequently, participants assessed their perceived control over the circle's color, based on their key-presses, via a rating scale that ranged from 0 % (no control) to 100 % (complete control). Contrary to Linser and Goschke (2007)'s findings, this experiment demonstrated that predictive information influenced the experience of agency only when primes were consciously processed. In experiment 2, utilizing symbolic (arrow) primes, N = 35 participants had to rate their feeling of control over the effect-stimulus' identity during a two-choice identification paradigm (i.e., they were instructed to press a key corresponding to a target stimulus; with a contingency between target and effect stimulus of 75 %/25 %). The results revealed no significant influence of subliminal priming on agency perceptions. In summary, this study implies that unconscious stimuli may not exert a substantial influence on the conscious experience of agency, underscoring the need for careful consideration of methodological aspects and experimental design's impact on observed phenomena.

Virtual occlusion effects on the perception of self-initiated visual stimuli.

Virtual reality (VR) has established itself as a useful tool in the study of human perception in the laboratory. A recent study introduced a new approach to examine visual sensory attenuation (SA) effects in VR. Hand movements triggered the appearance of Gabor stimuli, which were either presented behind the participant's hand - not rendered in VR ("virtual occlusion") - or elsewhere on the display. Virtual occlusion led to a rightward shift of the psychometric curve, suggesting that self-generated hand movements reduced the perceived contrast of the stimulus. Since such attenuation effects might provide a window into the predictive processing of the sensory and cognitive apparatus, we sought to better understand the nature of the virtual occlusion effects. In our study, the presentation of test stimuli was either self-initiated, self-initiated with a variable delay, or triggered externally; the test stimuli were occluded or not. In conflict with our hypothesis, we found moderate to strong evidence for an absence of any horizontal shifts between the psychometric curves. However, virtual occlusion was associated with a decrease in the slope of the psychometric function. Our results suggest that virtual occlusion attenuated the relative perceptual sensitivity, so that participants had more difficulty discriminating contrast differences when the test stimulus was presented behind the hand. We tentatively conclude that, in the visual domain, the discriminability of stimulus intensity is modified by internal predictive cues (i.e., proprioception), possibly linked to shifts in covert spatial attention.

Musicality as a predictive process.

Savage et al. argue for musicality as having evolved for the overarching purpose of social bonding. By way of contrast, we highlight contemporary predictive processing models of human cognitive functioning in which the production and enjoyment of music follows directly from the principle of prediction error minimization.

The attentional blink unveils the interplay between conscious perception, spatial attention and working memory encoding.

Our ability to perceive two events in close temporal succession is severely limited, a phenomenon known as the attentional blink. While the blink has served as a popular tool to prevent conscious perception, there is less research on its causes, and in particular on the role of conscious perception of the first event in triggering it. In three experiments, we disentangled the roles of spatial attention, conscious perception and working memory (WM) in causing the blink. We show that while allocating spatial attention to T1 is neither necessary nor sufficient for eliciting a blink, consciously perceiving it is necessary but not sufficient. When T1 was task irrelevant, consciously perceiving it triggered a blink only when it matched the attentional set for T2. We conclude that consciously perceiving a task-relevant event causes the blink, possibly because it triggers encoding of this event into WM. We discuss the implications of these findings for the relationship between spatial attention, conscious perception and WM, as well as for the distinction between access and phenomenal consciousness.

Unconscious arithmetic: Assessing the robustness of the results reported by Karpinski, Briggs, and Yale (2018).

In 2012, a study by Sklar et al. reported that participants could solve invisible subtractions. This notion of unconscious arithmetic has been influential because it challenges current theories of consciousness. In 2016, Karpinski et al. published a direct replication reporting evidence for unconscious addition rather than subtraction. About a year later, the study was retracted due to a computation error in the analysis pipeline. After this error was corrected, no evidence for unconscious addition nor subtraction was obtained. Recently, Karpinski et al. republished the study by applying the exclusion criteria used in Sklar et al. The reanalysis found weak evidence for unconscious subtraction. To assess the robustness of these results, we examine how sensitive the results are to data analytic decisions. We outline a set of 250 analyses that we consider justified to perform. We show that none of the analyses indicates evidence for unconscious subtraction.

A predictive coding account of bistable perception - a model-based fMRI study.

In bistable vision, subjective perception wavers between two interpretations of a constant ambiguous stimulus. This dissociation between conscious perception and sensory stimulation has motivated various empirical studies on the neural correlates of bistable perception, but the neurocomputational mechanism behind endogenous perceptual transitions has remained elusive. Here, we recurred to a generic Bayesian framework of predictive coding and devised a model that casts endogenous perceptual transitions as a consequence of prediction errors emerging from residual evidence for the suppressed percept. Data simulations revealed close similarities between the model's predictions and key temporal characteristics of perceptual bistability, indicating that the model was able to reproduce bistable perception. Fitting the predictive coding model to behavioural data from an fMRI-experiment on bistable perception, we found a correlation across participants between the model parameter encoding perceptual stabilization and the behaviourally measured frequency of perceptual transitions, corroborating that the model successfully accounted for participants' perception. Formal model comparison with established models of bistable perception based on mutual inhibition and adaptation, noise or a combination of adaptation and noise was used for the validation of the predictive coding model against the established models. Most importantly, model-based analyses of the fMRI data revealed that prediction error time-courses derived from the predictive coding model correlated with neural signal time-courses in bilateral inferior frontal gyri and anterior insulae. Voxel-wise model selection indicated a superiority of the predictive coding model over conventional analysis approaches in explaining neural activity in these frontal areas, suggesting that frontal cortex encodes prediction errors that mediate endogenous perceptual transitions in bistable perception. Taken together, our current work provides a theoretical framework that allows for the analysis of behavioural and neural data using a predictive coding perspective on bistable perception. In this, our approach posits a crucial role of prediction error signalling for the resolution of perceptual ambiguities.

Priming in a shape task but not in a category task under continuous flash suppression.

Continuous flash suppression (CFS) is an interocular suppression technique that uses high-contrast masks flashed to one eye to prevent conscious perception of images shown to the other eye. It has become widely used due to its strength and prolonged duration of suppression and its nearly deterministic control of suppression onset and offset. Recently, it has been proposed that action-relevant visual processing ascribed to the dorsal stream remains functional, while processing in the ventral stream is completely suppressed, when stimuli are invisible under CFS. Here we tested the hypothesis that the potentially dorsal-stream-based analysis of prime-stimulus elongation during CFS affects the categorization of manipulable target objects. In two behavioral experiments, we found evidence for priming in a shape task, but none for priming in a category task, when prime stimuli were rendered invisible using CFS. Our results thus support the notion that the representation of CF-suppressed stimuli is more limited than previously thought.

Investigating category- and shape-selective neural processing in ventral and dorsal visual stream under interocular suppression.

Recent behavioral and neuroimaging studies using continuous flash suppression (CFS) have suggested that action-related processing in the dorsal visual stream might be independent of perceptual awareness, in line with the "vision-for-perception" versus "vision-for-action" distinction of the influential dual-stream theory. It remains controversial if evidence suggesting exclusive dorsal stream processing of tool stimuli under CFS can be explained by their elongated shape alone or by action-relevant category representations in dorsal visual cortex. To approach this question, we investigated category- and shape-selective functional magnetic resonance imaging-blood-oxygen level-dependent responses in both visual streams using images of faces and tools. Multivariate pattern analysis showed enhanced decoding of elongated relative to non-elongated tools, both in the ventral and dorsal visual stream. The second aim of our study was to investigate whether the depth of interocular suppression might differentially affect processing in dorsal and ventral areas. However, parametric modulation of suppression depth by varying the CFS mask contrast did not yield any evidence for differential modulation of category-selective activity. Together, our data provide evidence for shape-selective processing under CFS in both dorsal and ventral stream areas and, therefore, do not support the notion that dorsal "vision-for-action" processing is exclusively preserved under interocular suppression.

Weighing the evidence for a dorsal processing bias under continuous flash suppression.

With the introduction of continuous flash suppression (CFS) as a method to render stimuli invisible and study unconscious visual processing, a novel hypothesis has gained popularity. It states that processes typically ascribed to the dorsal visual stream can escape CFS and remain functional, while ventral stream processes are suppressed when stimuli are invisible under CFS. This notion of a CFS-specific "dorsal processing bias" has been argued to be in line with core characteristics of the influential dual-stream hypothesis of visual processing which proposes a dissociation between dorsally mediated vision-for-action and ventrally mediated vision-for-perception. Here, we provide an overview of neuroimaging and behavioral studies that either examine this dorsal processing bias or base their conclusions on it. We show that both evidence for preserved ventral processing as well as lack of dorsal processing can be found in studies using CFS. To reconcile the diverging results, differences in the paradigms and their effects are worthy of future research. We conclude that given the current level of information a dorsal processing bias under CFS cannot be universally assumed.

Frontoparietal cortex mediates perceptual transitions in bistable perception.

During bistable vision, perception oscillates between two mutually exclusive percepts despite constant sensory input. Greater BOLD responses in frontoparietal cortex have been shown to be associated with endogenous perceptual transitions compared with "replay" transitions designed to closely match bistability in both perceptual quality and timing. It has remained controversial, however, whether this enhanced activity reflects causal influences of these regions on processing at the sensory level or, alternatively, an effect of stimulus differences that result in, for example, longer durations of perceptual transitions in bistable perception compared with replay conditions. Using a rotating Lissajous figure in an fMRI experiment on 15 human participants, we controlled for potential confounds of differences in transition duration and confirmed previous findings of greater activity in frontoparietal areas for transitions during bistable perception. In addition, we applied dynamic causal modeling to identify the neural model that best explains the observed BOLD signals in terms of effective connectivity. We found that enhanced activity for perceptual transitions is associated with a modulation of top-down connectivity from frontal to visual cortex, thus arguing for a crucial role of frontoparietal cortex in perceptual transitions during bistable perception.

Self-initiation enhances perceptual processing of auditory stimuli in an online study.

Understanding how the brain incorporates sensory and motor information will enable better theory building on human perception and behavior. In this study, we aimed to estimate the influence of predictive mechanisms on the magnitude and variability of sensory attenuation in two online samples. After the presentation of a visual cue stimulus, participants (Experiment 1: N = 224, Experiment 2: N = 84) compared the loudness of two consecutive tones in a two-alternative forced-choice task. In Experiment 1, the first tone was either self-initiated or not; in Experiment 2, the second tone was either self-initiated or not (active and passive condition, respectively). We further manipulated identity prediction (i.e., the congruence of pre-learned cue-sound combinations; congruent vs. incongruent), and the duration of the onset delay (to account for effects of attentional differences between the passive and active condition, 50 ms vs. 0 ms). We critically discuss our results within the framework of both classical (i.e., motor-based forward models) and contemporary approaches (i.e., predictive processing framework). Contrary to our preregistered hypothesis, we observed enhanced perceptual processing, instead of attenuation, for self-initiated auditory sensory input. Further, our results reveal an effect of fixed sound delays on the processing of motor and non-motor-based predictive information, and may point to according shifts in attention, leading to a perceptual bias. These results might best be captured by a hybrid explanatory model, combining predictions based on self-initiated motor action with a global predictive mechanism.

Temporal tuning properties along the human ventral visual stream.

Both our environment and our behavior contain many spatiotemporal regularities. Preferential and differential tuning of neural populations to these regularities can be demonstrated by assessing rate dependence of neural responses evoked during continuous periodic stimulation. Here, we used functional magnetic resonance imaging to measure regional variations of temporal sensitivity along the human ventral visual stream. By alternating one face and one house stimulus, we combined sufficient low-level signal modulation with changes in semantic meaning and could therefore drive all tiers of visual cortex strongly enough to assess rate dependence. We found several dissociations between early visual cortex and middle- and higher-tier regions. First, there was a progressive slowing down of stimulation rates yielding peak responses along the ventral visual stream. This finding shows the width of temporal integration windows to increase at higher hierarchical levels. Next, for fixed rates, early but not higher visual cortex responses additionally depended on the length of stimulus exposure, which may indicate increased persistence of responses to short stimuli at higher hierarchical levels. Finally, attention, which was recruited by an incidental task, interacted with stimulation rate and shifted tuning peaks toward lower frequencies. Together, these findings quantify neural response properties that are likely to be operational during natural vision and that provide putative neurofunctional substrates of mechanisms that are relevant in several psychophysical phenomena as masking and the attentional blink. Moreover, they illustrate temporal constraints for translating the deployment of attention into enhanced neural responses and thereby account for lower limits of attentional dwell time.

Posing for awareness: proprioception modulates access to visual consciousness in a continuous flash suppression task.

The rules governing the selection of which sensory information reaches consciousness are yet unknown. Of our senses, vision is often considered to be the dominant sense, and the effects of bodily senses, such as proprioception, on visual consciousness are frequently overlooked. Here, we demonstrate that the position of the body influences visual consciousness. We induced perceptual suppression by using continuous flash suppression. Participants had to judge the orientation a target stimulus embedded in a task-irrelevant picture of a hand. The picture of the hand could either be congruent or incongruent with the participants' actual hand position. When the viewed and the real hand positions were congruent, perceptual suppression was broken more rapidly than during incongruent trials. Our findings provide the first evidence of a proprioceptive bias in visual consciousness, suggesting that proprioception not only influences the perception of one's own body and self-consciousness, but also visual consciousness.

No conclusive evidence for number-induced attentional shifts in a temporal order judgement task.

The Spatial Numerical Association of Response Codes (SNARC) effect refers to the observation that relatively small (e.g., 1) and large numbers (e.g., 9) elicit faster left- and right-sided manual responses, respectively. In a variation known as the attentional SNARC effect, merely looking at numbers caused a left- or right-ward shift in covert spatial attention, depending on the number's magnitude. In our study, we probed the notion that numbers induce shifts of spatial attention in accordance with their position on a mental number line (MNL). Critically, we removed any putative spatial response code that may contaminate the responses. We used a square and a tilted square as targets, thereby situating the decisive response dimension in the ventral, non-spatial processing stream. In two experiments where numbers were used as non-informative cues preceding a temporal order judgement (TOJ) task, we did not observe a deflection of the locus of spatial attention as a function of the numerical magnitude of the cue. In a third experiment, finding a significant modulation of TOJ performance as a function of the pointing direction of arrow cues allowed us to rule out the possibility that the absence of any significant modulation in Experiments 1 and 2 was due to a lack of sensitivity of our task set-up. We conclude from the current findings that the spatial codes that the perception and naming of numbers potentially elicit are not in and by themselves sufficient to elicit deflections of spatial attention.

No effect of attentional modulation by spatial cueing in a masked numerical priming paradigm using continuous flash suppression (CFS).

One notion emerging from studies on unconscious visual processing is that different "blinding techniques" seem to suppress the conscious perception of stimuli at different levels of the neurocognitive architecture. However, even when only the results from a single suppression method are compared, the picture of the scope and limits of unconscious visual processing remains strikingly heterogeneous, as in the case of continuous flash suppression (CFS). To resolve this issue, it has been suggested that high-level semantic processing under CFS is facilitated whenever interocular suppression is attenuated by the removal of visuospatial attention. In this behavioral study, we aimed to further investigate this "CFS-attenuation-by-inattention" hypothesis in a numerical priming study using spatial cueing. Participants performed a number comparison task on a visible target number ("compare number to five"). Prime-target pairs were either congruent (both numbers smaller, or both larger than five) or incongruent. Based on the "CFS-attenuation-by-inattention" hypothesis, we predicted that reaction times (RTs) for congruent prime-target pairs should be faster than for incongruent ones, but only when the prime was presented at the uncued location. In the invisible condition, we observed no priming effects and thus no evidence in support of the "CFS-attenuation-by-inattention" hypothesis. In the visible condition, we found an inverse effect of prime-target congruency. Our results agree with the notion that the representation of CF-suppressed stimuli is fractionated, and limited to their basic, elemental features, thus precluding semantic processing.

Overlapping but separate number representations in the intraparietal sulcus-Probing format- and modality-independence in sighted Braille readers.

The Triple-Code Model stipulates that numerical information from different formats and modalities converges on a common magnitude representation in the Intraparietal Sulcus (IPS). To what extent the representations of all numerosity forms overlap remains unsolved. It has been postulated that the representation of symbolic numerosities (for example, Arabic digits) is sparser and grounded in an existing representation that codes for non-symbolic numerosity information (i.e., sets of objects). Other theories argue that numerical symbols represent a separate number category that emerges only during education. Here, we tested a unique group of sighted tactile Braille readers with numerosities 2, 4, 6 and 8 in three number notations: Arabic digits, sets of dots, tactile Braille numbers. Using univariate methods, we showed a consistent overlap in activations evoked by these three number notations. This result shows that all three used notations are represented in the IPS, which may suggest at least a partial overlap between the representations of the three notations used in this experiment. Using MVPA, we found that only non-automatized number information (Braille and sets of dots) allowed successful number classification. However, the numerosity of one notation could not be predicted above chance from the brain activation patterns evoked by another notation (no cross-classification). These results show that the IPS may host independent number codes in overlapping cortical circuits. In addition, they suggest that the level of training in encoding a given type of number information is an important factor that determines the amount of exploitable information and needs to be controlled for in order to identify the neural code underlying numerical information per se.

Differential BOLD activity associated with subjective and objective reports during "blindsight" in normal observers.

The study of conscious visual perception invariably necessitates some means of report. Report can be either subjective, i.e., an introspective evaluation of conscious experience, or objective, i.e., a forced-choice discrimination regarding different stimulus states. However, the link between report type and fMRI-BOLD signals has remained unknown. Here we used continuous flash suppression to render target images invisible, and observed a long-lasting dissociation between subjective report of visibility and human subjects' forced-choice localization of targets ("blindsight"). Our results show a robust dissociation between brain regions and type of report. We find subjective visibility effects in high-order visual areas even under equal objective performance. No significant BOLD difference was found between correct and incorrect trials in these areas when subjective report was constant. On the other hand, objective performance was linked to the accuracy of multivariate pattern classification mainly in early visual areas. Together, our data support the notion that subjective and objective reports tap cortical signals of different location and amplitude within the visual cortex.

Closing the gates to consciousness: distractors activate a central inhibition process.

The paradigm of distractor-induced blindness has previously been used to track the transition from unconscious to conscious visual processing. In a variation of this paradigm used in this study, participants (n = 13) had to detect an orientation change of tilted bars (target) embedded in a dynamic random pattern; the onset of the target was signaled by the presentation of a color cue. Occasional orientation changes preceding the cue served as distractors and severely impaired the target's detection. ERPs showed that a frontal negativity was cumulatively activated by the distractors, and early sensory components were not affected. In a control condition, the target was defined by a coherent motion of the bars. Orientation changes preceding the motion target did not affect its detection, and the frontal suppression process was not observed. However, we obtained a significant reduction of the sensory components. The data support the notion that distractors that share the target's features trigger a cumulative inhibition process preventing the conscious representation of the inhibited features. Explorative source modeling suggests that this process originates in the pFC. A top-down modulation of sensory processing could not be observed.

Give it time: neural evidence for distorted time perception and enhanced memory encoding in emotional situations.

Time perception is compromised in emotional situations, yet our ability to remember these events is enhanced. Here we suggest how the two phenomena might be functionally linked and describe the neural networks that underlie this association. We found that participants perceived an emotionally aversive stimulus longer than it was, compared to an immediately following neutral stimulus. These time estimation errors were in the same trials associated with better recognition memory for the emotionally aversive stimuli and poorer memory for the neutral stimuli. Functional imaging revealed that the superior frontal gyrus was activated during time perception with aversive stimuli, and the amygdala, putamen and insula showed activations that are specific to time estimation errors in this aversive context. We further found that activity in the insula and putamen was correlated with memory performance but only during over-estimation of time with the aversive stimuli. We suggest that processing is accelerated during the experience of emotionally aversive events, presumably in the service of memory-related operations, resulting in better encoding but at the expense of time perception accuracy.