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.

Reward Processing in Alcohol-Dependent Patients and First-Degree Relatives: Functional Brain Activity During Anticipation of Monetary Gains and Losses.

BACKGROUND: According to the reward deficiency syndrome and allostatic hypotheses, hyposensitivity of mesocorticolimbic regions to non-alcohol-related stimuli predisposes to dependence or is long-lastingly enhanced by chronic substance use. To date, no study has directly compared mesocorticolimbic brain activity during non-drug reward anticipation between alcohol-dependent, at risk, and healthy subjects. METHODS: Seventy-five abstinent alcohol-dependent human subjects (mean abstinence duration 957.66 days), 62 healthy first-degree relatives of alcohol-dependent individuals, and 76 healthy control subjects without family history of alcohol dependence performed a monetary incentive delay task. Functional magnetic resonance imaging data of the anticipation phase were analyzed, during which visual cues predicted that fast response to a target would result in monetary gain, avoidance of monetary loss, or a neutral outcome. RESULTS: During gain anticipation, there were no significant group differences. During loss anticipation, abstinent alcohol-dependent subjects showed lower activity in the left anterior insula compared with healthy control subjects without family history of alcohol dependence only (Montreal Neurological Institute [MNI] -25 19 -5; t206 = 4.17, familywise error corrected p = .009). However, this effect was no longer significant when age was included as a covariate. There were no group differences between abstinent alcohol-dependent subjects and healthy first-degree relatives or between healthy first-degree relatives and healthy control subjects during loss anticipation, respectively. CONCLUSIONS: Neither the neural reward deficiency syndrome nor the allostatic hypotheses are supported by the results. Future studies should investigate whether the incentive salience hypothesis allows for more accurate predictions regarding mesocorticolimbic brain activity of subjects with alcohol dependence and healthy individuals during reward and loss anticipation and further examine the neural substrates underlying a predisposition to dependence.

Pavlovian-to-Instrumental Transfer across Mental Disorders: A Review.

A mechanism known as Pavlovian-to-instrumental transfer (PIT) describes a phenomenon by which the values of environmental cues acquired through Pavlovian conditioning can motivate instrumental behavior. PIT may be one basic mechanism of action control that can characterize mental disorders on a dimensional level beyond current classification systems. Therefore, we review human PIT studies investigating subclinical and clinical mental syndromes. The literature prevails an inhomogeneous picture concerning PIT. While enhanced PIT effects seem to be present in non-substance-related disorders, overweight people, and most studies with AUD patients, no altered PIT effects were reported in tobacco use disorder and obesity. Regarding AUD and relapsing alcohol-dependent patients, there is mixed evidence of enhanced or no PIT effects. Additionally, there is evidence for aberrant corticostriatal activation and genetic risk, e.g., in association with high-risk alcohol consumption and relapse after alcohol detoxification. In patients with anorexia nervosa, stronger PIT effects elicited by low caloric stimuli were associated with increased disease severity. In patients with depression, enhanced aversive PIT effects and a loss of action-specificity associated with poorer treatment outcomes were reported. Schizophrenic patients showed disrupted specific but intact general PIT effects. Patients with chronic back pain showed reduced PIT effects. We provide possible reasons to understand heterogeneity in PIT effects within and across mental disorders. Further, we strengthen the importance of reliable experimental tasks and provide test-retest data of a PIT task showing moderate to good reliability. Finally, we point toward stress as a possible underlying factor that may explain stronger PIT effects in mental disorders, as there is some evidence that stress per se interacts with the impact of environmental cues on behavior by selectively increasing cue-triggered wanting. To conclude, we discuss the results of the literature review in the light of Research Domain Criteria, suggesting future studies that comprehensively assess PIT across psychopathological dimensions.

Probing the attentional modulation of unconscious processing under interocular suppression in a spatial cueing paradigm.

The debate about the scope and limits of unconscious visual processing under continuous flash suppression (CFS) has created a heterogeneous set of divergent findings that are yet to be reconciled. Attention has been suggested as an important factor in modulating the processing of suppressed visual information under CFS. Specifically, Eo et al. (2016) reported that semantic processing under CFS can be significantly facilitated when spatial attention is diverted away from the suppressed stimulus. Based on event-related potential (ERP) findings involving the N400, they proposed that inattention attenuates interocular suppression and thereby makes semantic processing available unconsciously, potentially reconciling conflicting evidence in the literature. In this study, we aimed to further investigate the "CFS-attenuation-by-inattention" hypothesis using functional magnetic resonance imaging (fMRI) and multivariate pattern analysis (MVPA). We tested whether the decodability of object category increases under CFS when attention is diverted away from the suppressed stimulus in a spatial cueing task. Our results provide no evidence for the "CFS-attenuation-by-inattention" hypothesis, but show higher decoding accuracies for visible stimuli than for invisible stimuli. We discuss the implications of our findings for the important endeavor of trying to reconcile the divergent reports of unconscious processing under CFS.

The Pervasive Problem of Post Hoc Data Selection in Studies on Unconscious Processing.

Studies on unconscious mental processes typically require that participants are unaware of some information (e.g., a visual stimulus). An important methodological question in this field of research is how to deal with data from participants who become aware of the critical stimulus according to some measure of awareness. While it has previously been argued that the post hoc selection of participants dependent on an awareness measure may often result in regression-to-the-mean artifacts (Shanks, 2017), a recent article (Sklar et al., 2021) challenged this conclusion claiming that the consideration of this statistical artifact might lead to unjustified rejections of true unconscious influences. In this reply, we explain this pervasive statistical problem with a basic and concrete example, show that Sklar et al. fundamentally mischaracterize it, and then refute the argument that the influence of the artifact has previously been overestimated. We conclude that, without safeguards, the method of post hoc data selection should never be employed in studies on unconscious processing.

Associations with monetary values do not influence access to awareness for faces.

Human faces can convey socially relevant information in various ways. Since the early detection of such information is crucial in social contexts, socially meaningful information might also have privileged access to awareness. This is indeed suggested by previous research using faces with emotional expressions. However, the social relevance of emotional faces is confounded with their physical stimulus characteristics. Here, we sought to overcome this problem by manipulating the relevance of face stimuli through classical conditioning: Participants had to learn the association between different face exemplars and high or low amounts of positive and negative monetary outcomes. Before and after the conditioning procedure, the time these faces needed to enter awareness was probed using continuous flash suppression, a variant of binocular rivalry. While participants successfully learned the association between the face stimuli and the respective monetary outcomes, faces with a high monetary value did not enter visual awareness faster than faces with a low monetary value after conditioning, neither for rewarding nor for aversive outcomes. Our results tentatively suggest that behaviorally relevant faces do not have privileged access to awareness when the assessment of the faces' relevance is dependent on the processing of face identity, as this requires complex stimulus processing that is likely limited at pre-conscious stages.

Unreliable feedback deteriorates information processing in primary visual cortex.

It is well-established that increased sensory uncertainty impairs perceptual decision-making and leads to degraded neural stimulus representations. Recently, we also showed that providing unreliable feedback to choices leads to changes in perceptual decision-making similar to those of increased stimulus noise: A deterioration in objective task performance, a decrease in subjective confidence and a lower reliance on sensory information for perceptual inference. To investigate the neural basis of such feedback-based changes in perceptual decision-making, in the present study, two groups of healthy human participants (n = 15 each) performed a challenging visual orientation discrimination task while undergoing functional magnetic resonance imaging (fMRI). Critically, one group received reliable feedback regarding their task performance in an intervention phase, whereas the other group correspondingly received unreliable feedback - thereby keeping stimulus information constant. The effects of feedback reliability on performance and stimulus representation in the primary visual cortex (V1) were studied by comparing the pre- and post-intervention test phases between the groups. Compared to participants who received reliable feedback, those receiving unreliable feedback showed a decline in task performance that was paralleled by reduced distinctness of fMRI response patterns in V1. These results show that environmental uncertainty can affect perceptual inference at the earliest cortical processing stages.

Pupil dilation as an implicit measure of appetitive Pavlovian learning.

Appetitive Pavlovian conditioning is a learning mechanism of fundamental biological and pathophysiological significance. Nonetheless, its exploration in humans remains sparse, which is partly attributed to the lack of an established psychophysiological parameter that aptly represents conditioned responding. This study evaluated pupil diameter and other ocular response measures (gaze dwelling time, blink duration and count) as indices of conditioning. Additionally, a learning model was used to infer participants' learning progress on the basis of their pupil dilation. Twenty-nine healthy volunteers completed an appetitive differential delay conditioning paradigm with a primary reward, while the ocular response measures along with other psychophysiological (heart rate, electrodermal activity, postauricular and eyeblink reflex) and behavioral (ratings, contingency awareness) parameters were obtained to examine the relation among different measures. A significantly stronger increase in pupil diameter, longer gaze duration and shorter eyeblink duration was observed in response to the reward-predicting cue compared to the control cue. The Pearce-Hall attention model best predicted the trial-by-trial pupil diameter. This conditioned response was corroborated by a pronounced heart rate deceleration to the reward-predicting cue, while no conditioning effect was observed in the electrodermal activity or startle responses. There was no discernible correlation between the psychophysiological response measures. These results highlight the potential value of ocular response measures as sensitive indices for representing appetitive conditioning.

The influence of gaze direction on food preferences.

In our information-rich environment, the gaze direction of another indicates their current focus of attention. Following the gaze of another, results in gaze-evoked shifts in joint attention, a phenomenon critical for the functioning of social cognition. Previous research in joint attention has shown that objects that are attended by another are more liked than ignored objects. Here, we investigated this effect of gaze-cueing on participants' preferences for unknown food items. Participants provided their willingness to pay (WTP), taste and health preferences for food items before and after a standard gaze-cueing paradigm. We observed a significant effect of gaze-cueing on participants' WTP bids. Specifically, participants were willing to pay more money for the food items that were looked at by another person. In contrast, there was a decrease in preference for the food items that were ignored by another person. Interestingly, this increase in WTP occurred without participants' awareness of the contingency between the cue and target. These results highlight the influence of social information on human choice behavior and lay the foundation for experiments in neuromarketing and consumer decision making.

Access to awareness of direct gaze is related to autistic traits.

BACKGROUND: The atypical processing of eye contact is a characteristic hallmark of autism spectrum disorder (ASD). The severity of these symptoms, however, is thought to lie on a continuum that extends into the typical population. While behavioural evidence shows that differences in social cognitive tasks in typically developed (TD) adults are related to the levels of autistic-like traits, it remains unknown whether such a relation exists for the sensitivity to direct gaze. METHODS: In two experiments, we measured reaction times to detect the faces with direct and averted gaze, suppressed from awareness, i.e. the access to awareness. In experiment 1, we tested N = 19 clinically diagnosed adults with ASD and N = 22 TD matched controls, while in experiment 2, we tested an independent sample of N = 20 TD adults. RESULTS: In line with the literature, experiment 1 showed preferential processing of direct gaze in the TD group but not in the ASD group. Importantly, we found a linear relationship in both experiments between the levels of autistic traits within the groups of TD participants and their sensitivity to direct gaze: with increasing autistic characteristics, there was a decrease in sensitivity to direct gaze. CONCLUSION: These results provide the first evidence that differences in gaze processing and the sensitivity to direct gaze are already present in individuals with subclinical levels of autistic traits. Furthermore, they lend support to the continuum view of the disorder and could potentially help in an earlier diagnosis of individuals at high risk for autism.

Investigating masked priming along the "vision-for-perception" and "vision-for-action" dimensions of unconscious processing.

The study of nonconscious priming is rooted in a long research tradition in experimental psychology and plays an important role for a range of topics, including visual recognition, emotion, decision making, and memory. Prime stimuli can be transiently suppressed from awareness by using a variety of psychophysical paradigms. The aim is to understand which stimulus features can be processed nonconsciously and influence behavior toward subsequently presented probe stimuli. Here, we tested the notion that continuous flash suppression (CFS), a relatively new method of interocular suppression, selectively disrupts stimulus identification mediated by the ventral "vision-for-perception" pathway, while preserving action-relevant stimulus features processed by the dorsal "vision-for-action" pathway. Given the far-reaching implications of this notion for the influential two visual systems hypothesis, and visual cognition in general, we investigated its empirical basis in a series of seven masked priming experiments using CFS. We did not find evidence for nonconscious priming of object categorization by action-relevant features. Based on these results, we recommend skepticism about the notion that the processing of action-relevant features under CFS is selectively preserved in the "vision-for-action" pathway. Second, we conclude that CFS experiments are less informative than approaches using visible stimuli, when the aim is to gather data in relation to the two visual systems hypothesis. Third, we propose that future nonconscious priming studies should carefully consider the position of suppression paradigms within a functional hierarchy of unconscious processing, thus constraining hypothesis generation to effects that are plausible given the employed methodology. (PsycINFO Database Record (c) 2018 APA, all rights reserved).

Unconscious avoidance of eye contact in autism spectrum disorder.

Atypical responses to direct gaze are one of the most characteristic hallmarks of autism spectrum disorder (ASD). The cause and mechanism underlying this phenomenon, however, have remained unknown. Here we investigated whether the atypical responses to eye gaze in autism spectrum disorder is dependent on the conscious perception of others' faces. Face stimuli with direct and averted gaze were rendered invisible by interocular suppression and eye movements were recorded from participants with ASD and an age and sex matched control group. Despite complete unawareness of the stimuli, the two groups differed significantly in their eye movements to the face stimuli. In contrast to the significant positive saccadic index observed in the TD group, indicating an unconscious preference to the face with direct gaze, the ASD group had no such preference towards direct gaze and instead showed a tendency to prefer the face with averted gaze, suggesting an unconscious avoidance of eye contact. These results provide the first evidence that the atypical response to eye contact in ASD is an unconscious and involuntary response. They provide a better understanding of the mechanism of gaze avoidance in autism and might lead to new diagnostic and therapeutic interventions.

What We Talk about When We Talk about Unconscious Processing - A Plea for Best Practices.

In this perspective article, we first outline the large diversity of methods, measures, statistical analyses, and concepts in the field of the experimental study of unconscious processing. We then suggest that this diversity implies that comparisons between different studies on unconscious processing are fairly limited, especially when stimulus awareness has been assessed in different ways. Furthermore, we argue that flexible choices of methods and measures will inevitably lead to an overestimation of unconscious processes. In the concluding paragraph, we briefly present solutions and strategies for future research. We make a plea for the introduction of "best practices," similar to previous attempts to constitute practicing standards for functional magnetic resonance imaging (fMRI) and electroencephalography (EEG).

Neural mechanisms of reinforcement learning in unmedicated patients with major depressive disorder.

According to current concepts, major depressive disorder is strongly related to dysfunctional neural processing of motivational information, entailing impairments in reinforcement learning. While computational modelling can reveal the precise nature of neural learning signals, it has not been used to study learning-related neural dysfunctions in unmedicated patients with major depressive disorder so far. We thus aimed at comparing the neural coding of reward and punishment prediction errors, representing indicators of neural learning-related processes, between unmedicated patients with major depressive disorder and healthy participants. To this end, a group of unmedicated patients with major depressive disorder (n = 28) and a group of age- and sex-matched healthy control participants (n = 30) completed an instrumental learning task involving monetary gains and losses during functional magnetic resonance imaging. The two groups did not differ in their learning performance. Patients and control participants showed the same level of prediction error-related activity in the ventral striatum and the anterior insula. In contrast, neural coding of reward prediction errors in the medial orbitofrontal cortex was reduced in patients. Moreover, neural reward prediction error signals in the medial orbitofrontal cortex and ventral striatum showed negative correlations with anhedonia severity. Using a standard instrumental learning paradigm we found no evidence for an overall impairment of reinforcement learning in medication-free patients with major depressive disorder. Importantly, however, the attenuated neural coding of reward in the medial orbitofrontal cortex and the relation between anhedonia and reduced reward prediction error-signalling in the medial orbitofrontal cortex and ventral striatum likely reflect an impairment in experiencing pleasure from rewarding events as a key mechanism of anhedonia in major depressive disorder.

Enhanced predictive signalling in schizophrenia.

Positive symptoms of schizophrenia such as delusions and hallucinations are thought to arise from an alteration in predictive mechanisms of the brain. Here, we empirically tested the hypothesis that schizophrenia is associated with an enhanced signalling of higher-level predictions that shape perception into conformity with acquired beliefs. Twenty-one patients with schizophrenia and twenty-eight healthy controls matched for age and gender took part in a functional magnetic resonance imaging (fMRI) experiment that assessed the effect of an experimental manipulation of cognitive beliefs on the perception of an ambiguous visual motion stimulus. At the behavioural level, there was a generally weaker effect of experimentally induced beliefs on perception in schizophrenia patients compared with controls, but a positive correlation between the effect of beliefs on perception and the severity of positive symptoms. At the neural level, belief-related connectivity between a region encoding beliefs in the orbitofrontal cortex (OFC) and a region encoding visual motion in the visual cortex (V5) was higher in patients compared with controls, indicating a stronger impact of cognitive beliefs on visual processing in schizophrenia. We suggest that schizophrenia might be associated with a generally weaker acquisition of externally generated beliefs and a compensatory increase in the effect of beliefs on sensory processing. Our current results are in line with the notion that enhanced signalling of higher-level predictions that shape perception into conformity with acquired beliefs might underlie positive symptoms in schizophrenia. Hum Brain Mapp 38:1767-1779, 2017. © 2017 Wiley Periodicals, Inc.

Probing the influence of unconscious fear-conditioned visual stimuli on eye movements.

Efficient threat detection from the environment is critical for survival. Accordingly, fear-conditioned stimuli receive prioritized processing and capture overt and covert attention. However, it is unknown whether eye movements are influenced by unconscious fear-conditioned stimuli. We performed a classical fear-conditioning procedure and subsequently recorded participants' eye movements while they were exposed to fear-conditioned stimuli that were rendered invisible using interocular suppression. Chance-level performance in a forced-choice-task demonstrated unawareness of the stimuli. Differential skin conductance responses and a change in participants' fearfulness ratings of the stimuli indicated the effectiveness of conditioning. However, eye movements were not biased towards the fear-conditioned stimulus. Preliminary evidence suggests a relation between the strength of conditioning and the saccadic bias to the fear-conditioned stimulus. Our findings provide no strong evidence for a saccadic bias towards unconscious fear-conditioned stimuli but tentative evidence suggests that such an effect may depend on the strength of the conditioned response.

Decomposing Self-Control: Individual Differences in Goal Pursuit Despite Interfering Aversion, Temptation, and Distraction.

Self-control can be defined as the ability to exert control over ones impulses. Currently, most research in the area relies on self-report. Focusing on attentional control processes involved in self-control, we modified a spatial selective attentional cueing task to test three domains of self-control experimentally in one task using aversive, tempting, and neutral picture-distractors. The aims of the study were (1) to investigate individual differences in the susceptibility to aversive, tempting, and neutral distraction within one paradigm and (2) to test the association of these three self-control domains to conventional measures of self-control including self-report. The final sample consisted of 116 participants. The task required participants to identify target letters "E" or "F" presented at a cued target location while the distractors were presented. Behavioral and eyetracking data were obtained during the performance of the task. High task performance was encouraged via monetary incentives. In addition to the attentional self-control task, self-reported self-control was assessed and participants performed a color Stroop task, an unsolvable anagram task and a delay of gratification task using chocolate sweets. We found that aversion, temptation, and neutral distraction were associated with significantly increased error rates, reaction times and gaze pattern deviations. Overall task performance on our task correlated with self-reported self-control ability. Measures of aversion, temptation, and distraction showed moderate split-half reliability, but did not correlate with each other across participants. Additionally, participants who made a self-controlled decision in the delay of gratification task were less distracted by temptations in our task than participants who made an impulsive choice. Our individual differences analyses suggest that (1) the ability to endure aversion, resist temptations and ignore neutral distractions are independent of each other and (2) these three domains are related to other measures of self-control.

Gaze Direction Modulates the Relation between Neural Responses to Faces and Visual Awareness.

Gaze direction and especially direct gaze is a powerful nonverbal cue that plays an important role in social interactions. Here we studied the neural mechanisms underlying the privileged access of direct gaze to visual awareness. We performed functional magnetic resonance imaging in healthy human volunteers who were exposed to faces with direct or averted gaze under continuous flash suppression, thereby manipulating their awareness of the faces. A gaze processing network comprising fusiform face area (FFA), superior temporal sulcus, amygdala, and intraparietal sulcus showed overall reduced neural responses when participants reported to be unaware of the faces. Interestingly, direct gaze elicited greater responses than averted gaze when participants were aware of the faces, but smaller responses when they were unaware. Additional between-subject correlation and single-trial analyses indicated that this pattern of results was due to a modulation of the relationship between neural responses and awareness by gaze direction: with increasing neural activation in the FFA, direct-gaze faces entered awareness more readily than averted-gaze faces. These findings suggest that for direct gaze, lower levels of neural activity are sufficient to give rise to awareness than for averted gaze, thus providing a neural basis for privileged access of direct gaze to awareness. Significance statement: Another person's eye gaze directed at oneself is a powerful social signal acting as a catalyst for further communication. Here, we studied the neural mechanisms underlying the prioritized access of direct gaze to visual awareness in healthy human volunteers and show that with increasing neural activation, direct-gaze faces enter awareness more readily than averted-gaze faces. This suggests that for a socially highly relevant cue like direct gaze, lower levels of neural activity are sufficient to give rise to awareness compared with averted gaze, possibly because the human brain is attuned to the efficient neural processing of direct gaze due to the biological importance of eye contact for social interactions.

Making eye contact without awareness.

Direct gaze is a potent non-verbal signal that establishes a communicative connection between two individuals, setting the course for further interactions. Although consciously perceived faces with direct gaze have been shown to capture attention, it is unknown whether an attentional preference for these socially meaningful stimuli exists even in the absence of awareness. In two experiments, we recorded participants' eye movements while they were exposed to faces with direct and averted gaze rendered invisible by interocular suppression. Participants' inability to correctly guess the occurrence of the faces in a manual forced-choice task demonstrated complete unawareness of the faces. However, eye movements were preferentially directed towards faces with direct compared to averted gaze, indicating a specific sensitivity to others' gaze directions even without awareness. This oculomotor preference suggests that a rapid and automatic establishment of mutual eye contact constitutes a biological advantage, which could be mediated by fast subcortical pathways in the human brain.