Extracting probability in the absence of visual awareness.

Extracting statistical regularities from the environment is crucial for survival. It allows us to learn cues for where and when future events will occur. Can we learn these associations even when the cues are not consciously perceived? Can these unconscious processes integrate information over long periods of time? We show that human visual system can track the probability of location contingency between an unconscious prime and a conscious target over a period of time of minutes. In a series of psychophysical experiments, we adopted an exogenous priming paradigm and manipulated the location contingency between a masked prime and a visible target (i.e., how likely the prime location predicted the target location). The prime's invisibility was verified both subjectively and objectively. Although the participants were unaware of both the existence of the prime and the prime-target contingency, our results showed that the probability of location contingency was tracked and manifested in the subsequent priming effect. When participants were first entrained into the fully predictive prime-target probability, they exhibited faster responses to the more predictive location. On the contrary, when no contingency existed between the prime and target initially, participants later showed faster responses to the less predictive location. These results were replicated in two more experiments with increased statistical power and a fine-grained delineation of prime awareness. Together, we report that the human visual system is capable of tracking unconscious probability over a period of time, demonstrating how implicit and uncertain regularity guides behavior.

Attractiveness in the Eyes: A Possibility of Positive Loop between Transient Pupil Constriction and Facial Attraction.

Contrary to the long-held belief of a close linkage between pupil dilation and attractiveness, we found an early and transient pupil constriction response when participants viewed an attractive face (and the effect of luminance/contrast was controlled). While human participants were making an attractiveness judgment on faces, their pupil constricted more for the more attractive (as-to-be-rated) faces. Further experiments showed that the effect of pupil constriction to attractiveness judgment extended to intrinsically esthetic visual objects such as natural scene images (as well as faces) but not to line-drawing geometric figures. When participants were asked to judge the roundness of faces, pupil constriction still correlated with their attractiveness but not the roundness rating score, indicating the automaticity of the pupil constriction to attractiveness. When pupillary responses were manipulated implicitly by relative background luminance changes (from the prestimulus screen), the facial attractiveness ratings were in accordance with the amount of pupil constriction, which could not be explained solely by simultaneous or sequential luminance contrast. The overall results suggest that pupil constriction not only reflects but, as a part of self-monitoring and attribution mechanisms, also possibly contributes to facial attractiveness implicitly.

Eye movement analysis with switching hidden Markov models.

Here we propose the eye movement analysis with switching hidden Markov model (EMSHMM) approach to analyzing eye movement data in cognitive tasks involving cognitive state changes. We used a switching hidden Markov model (SHMM) to capture a participant's cognitive state transitions during the task, with eye movement patterns during each cognitive state being summarized using a regular HMM. We applied EMSHMM to a face preference decision-making task with two pre-assumed cognitive states-exploration and preference-biased periods-and we discovered two common eye movement patterns through clustering the cognitive state transitions. One pattern showed both a later transition from the exploration to the preference-biased cognitive state and a stronger tendency to look at the preferred stimulus at the end, and was associated with higher decision inference accuracy at the end; the other pattern entered the preference-biased cognitive state earlier, leading to earlier above-chance inference accuracy in a trial but lower inference accuracy at the end. This finding was not revealed by any other method. As compared with our previous HMM method, which assumes no cognitive state change (i.e., EMHMM), EMSHMM captured eye movement behavior in the task better, resulting in higher decision inference accuracy. Thus, EMSHMM reveals and provides quantitative measures of individual differences in cognitive behavior/style, making a significant impact on the use of eyetracking to study cognitive behavior across disciplines.

Neural computations mediating one-shot learning in the human brain.

Incremental learning, in which new knowledge is acquired gradually through trial and error, can be distinguished from one-shot learning, in which the brain learns rapidly from only a single pairing of a stimulus and a consequence. Very little is known about how the brain transitions between these two fundamentally different forms of learning. Here we test a computational hypothesis that uncertainty about the causal relationship between a stimulus and an outcome induces rapid changes in the rate of learning, which in turn mediates the transition between incremental and one-shot learning. By using a novel behavioral task in combination with functional magnetic resonance imaging (fMRI) data from human volunteers, we found evidence implicating the ventrolateral prefrontal cortex and hippocampus in this process. The hippocampus was selectively "switched" on when one-shot learning was predicted to occur, while the ventrolateral prefrontal cortex was found to encode uncertainty about the causal association, exhibiting increased coupling with the hippocampus for high-learning rates, suggesting this region may act as a "switch," turning on and off one-shot learning as required.

Decision ambiguity is mediated by a late positive potential originating from cingulate cortex.

People often make decisions in the face of ambiguous information, but it remains unclear how ambiguity is represented in the brain. We used three types of ambiguous stimuli and combined EEG and fMRI to examine the neural representation of perceptual decisions under ambiguity. We identified a late positive potential, the LPP, which differentiated levels of ambiguity, and which was specifically associated with behavioral judgments about choices that were ambiguous, rather than passive perception of ambiguous stimuli. Mediation analyses together with two further control experiments confirmed that the LPP was generated only when decisions are made (not during mere perception of ambiguous stimuli), and only when those decisions involved choices on a dimension that is ambiguous. A further control experiment showed that a stronger LPP arose in the presence of ambiguous stimuli compared to when only unambiguous stimuli were present. Source modeling suggested that the LPP originated from multiple loci in cingulate cortex, a finding we further confirmed using fMRI and fMRI-guided ERP source prediction. Taken together, our findings argue for a role of an LPP originating from cingulate cortex in encoding decisions based on task-relevant perceptual ambiguity, a process that may in turn influence confidence judgment, response conflict, and error correction.

The effects of incentive framing on performance decrements for large monetary outcomes: behavioral and neural mechanisms.

There is a nuanced interplay between the provision of monetary incentives and behavioral performance. Individuals' performance typically increases with increasing incentives only up to a point, after which larger incentives may result in decreases in performance, a phenomenon known as "choking." We investigated the influence of incentive framing on choking effects in humans: in one condition, participants performed a skilled motor task to obtain potential monetary gains; in another, participants performed the same task to avoid losing a monetary amount. In both the gain and loss frame, the degree of participants' behavioral loss aversion was correlated with their susceptibility to choking effects. However, the effects were markedly different in the gain and loss frames: individuals with higher loss aversion were susceptible to choking for large prospective gains and not susceptible to choking for large prospective losses, whereas individuals with low loss aversion choked for large prospective losses but not for large prospective gains. Activity in the ventral striatum was predictive of performance decrements in both the gain and loss frames. Moreover, a mediation analysis revealed that behavioral loss aversion hindered performance via the influence of ventral striatal activity on motor performance. Our findings indicate that the framing of an incentive has a profound effect on an individual's susceptibility to choking effects, which is contingent on their loss aversion. Furthermore, we demonstrate that the ventral striatum serves as an interface between incentive-driven motivation and instrumental action, regardless of whether incentives are framed in terms of potential losses or gains.

The hand sees visual periphery better than the eye: motor-dependent visual motion analyses.

Information pertaining to visual motion is used in the brain not only for conscious perception but also for various kinds of motor controls. In contrast to the increasing amount of evidence supporting the dissociation of visual processing for action versus perception, it is less clear whether the analysis of visual input is shared for characterizing various motor outputs, which require different kinds of interactions with environments. Here we show that, in human visuomotor control, motion analysis for quick hand control is distinct from that for quick eye control in terms of spatiotemporal analysis and spatial integration. The amplitudes of implicit and quick hand and eye responses induced by visual motion stimuli differently varied with stimulus size and pattern smoothness (e.g., spatial frequency). Surprisingly, the hand response did not decrease even when the visual motion with a coarse pattern was mostly occluded over the visual center, whereas the eye response markedly decreased. Since these contrasts cannot be ascribed to any difference in motor dynamics, they clearly indicate different spatial integration of visual motion for the individual motor systems. Going against the overly unified hierarchical view of visual analysis, our data suggest that visual motion analyses are separately tailored from early levels to individual motor modalities. Namely, the hand and eyes see the external world differently.

Practice makes imperfect: stronger implicit interference with practice in individuals at high risk of developing Alzheimer's disease.

Early screening to determine patient risk of developing Alzheimer's will allow better interventions and planning but necessitates accessible methods such as behavioral biomarkers. Previously, we showed that cognitively healthy older individuals whose cerebrospinal fluid amyloid/tau ratio indicates high risk of cognitive decline experienced implicit interference during a high-effort task, signaling early changes in attention. To further investigate attention's effect on implicit interference, we analyzed two experiments completed sequentially by the same high- and low-risk individuals. We hypothesized that if attention modulates interference, practice would affect the influence of implicit distractors. Indeed, while both groups experienced a strong practice effect, the association between practice and interference effects diverged between groups: stronger practice effects correlated with more implicit interference in high-risk participants, but less interference in low-risk individuals. Furthermore, low-risk individuals showed a positive correlation between implicit interference and EEG low-range alpha event-related desynchronization when switching from high- to low-load tasks. This suggests that lower attention on the task was correlated with stronger interference, a typical phenomenon in the younger population. These results demonstrate how attention impacts implicit interference and highlight early differences in perception between high- and low-risk individuals.

Performance Dip in motor response induced by task-irrelevant weaker coherent visual motion signals.

The Performance Dip is a newly characterized behavioral phenomenon, where, paradoxically, a weaker task-irrelevant visual stimulus causes larger disturbances on the accuracy of a main letter identification task than a stronger stimulus does. Understanding mechanisms of the Performance Dip may provide insight into unconsciousness behavior. Here, we investigated the generalization of the Performance Dip. Specifically, we tested whether the Performance Dip occurs in a motion-related Simon task, and if so, whether the Performance Dip involves the same brain region, that is, the dorsolateral prefrontal cortex (DLPFC), previously implicated in the Performance Dip, or the supplementary motor area (SMA) and pre-SMA, implicated in a motion-related Simon Task. Subjects made manual directional responses according to the color of stochastic moving dots while ignoring the global direction of moving dots, which could be either congruent or incongruent to the response appropriate to the main task. We found that weak incongruent task-irrelevant stimuli caused a Performance Dip, in which the SMA and pre-SMA, rather than DLPFC, played critical roles. Our results suggest a possible common brain mechanism across different neural circuits, in which weak, but not strong, task-irrelevant information is free from inhibition and intrudes into neural circuits relevant to the main task.

Roles of familiarity and novelty in visual preference judgments are segregated across object categories.

Understanding preference decision making is a challenging problem because the underlying process is often implicit and dependent on context, including past experience. There is evidence for both familiarity and novelty as critical factors for preference in adults and infants. To resolve this puzzling contradiction, we examined the cumulative effects of visual exposure in different object categories, including faces, natural scenes, and geometric figures, in a two-alternative preference task. The results show a clear segregation of preference across object categories, with familiarity preference dominant in faces and novelty preference dominant in natural scenes. No strong bias was observed in geometric figures. The effects were replicated even when images were converted to line drawings, inverted, or presented only briefly, and also when spatial frequency and contour distribution were controlled. The effects of exposure were reset by a blank of 1 wk or 3 wk. Thus, the category-specific segregation of familiarity and novelty preferences is based on quick visual categorization and cannot be caused by the difference in low-level visual features between object categories. Instead, it could be due either to different biological significances/attractiveness criteria across these categories, or to some other factors, such as differences in within-category variance and adaptive tuning of the perceptual system.

Practice makes imperfect: stronger implicit interference with practice in individuals at high risk of developing Alzheimer's disease.

Early screening to determine patient risk of developing Alzheimer's will allow better interventions and planning but necessitates accessible methods such as behavioral biomarkers. Previously, we showed that cognitively healthy older individuals whose cerebrospinal fluid amyloid / tau ratio indicates high risk of cognitive decline experienced implicit interference during a high-effort task, signaling early changes in attention. To further investigate attention's effect on implicit interference, we analyzed two experiments completed sequentially by the same high- and low-risk individuals. We hypothesized that if attention modulates interference, practice would affect the influence of implicit distractors. Indeed, while both groups experienced a strong practice effect, the association between practice and interference effects diverged between groups: stronger practice effects correlated with more implicit interference in high-risk participants, but less interference in low-risk individuals. Furthermore, low-risk individuals showed a positive correlation between implicit interference and EEG low-range alpha event-related desynchronization when switching from high- to low-load tasks. These results demonstrate how attention impacts implicit interference and highlight early differences in cognition between high- and low-risk individuals.

The connection between heart rate variability (HRV), neurological health, and cognition: A literature review.

The heart and brain have bi-directional influences on each other, including autonomic regulation and hemodynamic connections. Heart rate variability (HRV) measures variation in beat-to-beat intervals. New findings about disorganized sinus rhythm (erratic rhythm, quantified as heart rate fragmentation, HRF) are discussed and suggest overestimation of autonomic activities in HRV changes, especially during aging or cardiovascular events. When excluding HRF, HRV is regulated via the central autonomic network (CAN). HRV acts as a proxy of autonomic activity and is associated with executive functions, decision-making, and emotional regulation in our health and wellbeing. Abnormal changes of HRV (e.g., decreased vagal functioning) are observed in various neurological conditions including mild cognitive impairments, dementia, mild traumatic brain injury, migraine, COVID-19, stroke, epilepsy, and psychological conditions (e.g., anxiety, stress, and schizophrenia). Efforts are needed to improve the dynamic and intriguing heart-brain interactions.

Orbitofrontal cortical activity during repeated free choice.

Neurons in the orbitofrontal cortex (OFC) have been shown to encode subjective values, suggesting a role in preference-based decision-making, although the precise relation to choice behavior is unclear. In a repeated two-choice task, subjective values of each choice can account for aggregate choice behavior, which is the overall likelihood of choosing one option over the other. Individual choices, however, are impossible to predict with knowledge of relative subjective values alone. In this study we investigated the role of internal factors in choice behavior with a simple but novel free-choice task and simultaneous recording from individual neurons in nonhuman primate OFC. We found that, first, the observed sequences of choice behavior included periods of exceptionally long runs of each of two available options and periods of frequent switching. Neither a satiety-based mechanism nor a random selection process could explain the observed choice behavior. Second, OFC neurons encode important features of the choice behavior. These features include activity selective for exceptionally long runs of a given choice (stay selectivity) as well as activity selective for switches between choices (switch selectivity). These results suggest that OFC neural activity, in addition to encoding subjective values on a long timescale that is sensitive to satiety, also encodes a signal that fluctuates on a shorter timescale and thereby reflects some of the statistically improbable aspects of free-choice behavior.

Shared visual attention reduces hindsight bias.

Hindsight bias is the tendency to retrospectively think of outcomes as being more foreseeable than they actually were. It is a robust judgment bias and is difficult to correct (or "debias"). In the experiments reported here, we used a visual paradigm in which performers decided whether blurred photos contained humans. Evaluators, who saw the photos unblurred and thus knew whether a human was present, estimated the proportion of participants who guessed whether a human was present. The evaluators exhibited visual hindsight bias in a way that matched earlier data from judgments of historical events surprisingly closely. Using eye tracking, we showed that a higher correlation between the gaze patterns of performers and evaluators (shared attention) is associated with lower hindsight bias. This association was validated by a causal method for debiasing: Showing the gaze patterns of the performers to the evaluators as they viewed the stimuli reduced the extent of hindsight bias.

Overlapping responses for the expectation of juice and money rewards in human ventromedial prefrontal cortex.

Although much is known about the neural substrates of reward, the question of whether expectation of different types of reinforcers engage distinct or overlapping brain circuitry has not been addressed definitively. In the present study, human subjects, while being scanned with functional magnetic resonance imaging, performed a simple reward-based action selection task to obtain different magnitudes of either monetary outcomes (winning or losing money) or juice outcomes (pleasant apple juice or an unpleasant salt flavor). At the group level, we found partially overlapping value-related activity within ventromedial prefrontal cortex (vmPFC) during anticipation of juice and money reward outcomes. Analogous results were found in the right anterior insula, except that this region showed negative correlations as a function of increasing expected reward. These results indicate that vmPFC and anterior insula contain overlapping representations of anticipatory value, consistent with the existence of a common currency for the value of expected outcomes in these regions.

Crossmodal Postdiction: Conscious Perception as Revisionist History.

Postdiction occurs when later stimuli influence the perception of earlier stimuli. As the multisensory science field has grown in recent decades, the investigation of crossmodal postdictive phenomena has also expanded. Crossmodal postdiction can be considered (in its simplest form) the phenomenon in which later stimuli in one modality influence earlier stimuli in another modality (e.g., Intermodal Apparent Motion). Crossmodal postdiction can also appear in more nuanced forms, such as unimodal postdictive illusions (e.g., Apparent Motion) that are influenced by concurrent crossmodal stimuli (e.g., Crossmodal Influence on Apparent Motion), or crossmodal illusions (e.g., the Double Flash Illusion) that are influenced postdictively by a stimulus in one or the other modality (e.g., a visual stimulus in the Illusory Audiovisual Rabbit Illusion). In this review, these and other varied forms of crossmodal postdiction will be discussed. Three neuropsychological models proposed for unimodal postdiction will be adapted to the unique aspects of processing and integrating multisensory stimuli. Crossmodal postdiction opens a new window into sensory integration, and could potentially be used to identify new mechanisms of crossmodal crosstalk in the brain.

Stronger implicit interference in cognitively healthy older participants with higher risk of Alzheimer's disease.

Introduction: Abnormal cerebrospinal fluid amyloid beta (Aß)42 and tau levels have been revealed decades before symptoms onset in Alzheimer's disease (AD); however, the examination is usually invasive and inaccessible to most people. We thus aimed to develop a non-invasive behavioral test that targets early potential cognitive changes to gauge cognitive decline. Specifically, we hypothesized that older cognitive healthy participants would exhibit comparable performance when the task was explicit and relied on conscious cognition. However, when the task was implicit, the performance of participants at high and low risks for AD would bifurcate. That is, early changes in unconscious cognition could be linked to cognitive health. Methods: We measured implicit interference elicited by an imperceptible distractor in cognitively healthy elderly participants with normal (low risk) and pathological (high risk) Aß42/total tau ratio. Participants were required to perform a Stroop task (word-naming or color-naming on an ink-semantics inconsistent word) with a visually masked distractor presented prior to the target task. Results: We found that, under a high-effort task (i.e., color-naming in the Stroop task), high-risk participants suffered interference when the imperceptible distractor and the subsequent target were incongruent in the responses they triggered. Their reaction times were slowed down by approximately 4%. This implicit interference was not found in the low-risk participants. Discussion: These findings indicate that weakened inhibition of distracting implicit information can be a potential behavioral biomarker of early identification of AD pathology. Our study thus offers a new experimental paradigm to reveal early pathological aging by assessing how individuals respond to subperceptual threshold visual stimuli.

An integration of color and motion information in visual scene analyses.

To analyze complex scenes efficiently, the human visual system performs perceptual groupings based on various features (e.g., color and motion) of the visual elements in a scene. Although previous studies demonstrated that such groupings can be based on a single feature (e.g., either color or motion information), here we show that the visual system also performs scene analyses based on a combination of two features. We presented subjects with a mixture of red and green dots moving in various directions. Although the pairings between color and motion information were variable across the dots (e.g., one red dot moved upward while another moved rightward), subjects' perceptions of the color-motion pairings were significantly biased when the randomly paired dots were flanked by additional dots with consistent color-motion pairings. These results indicate that the visual system resolves local ambiguities in color-motion pairings using unambiguous pairings in surrounds, demonstrating a new type of scene analysis based on the combination of two featural cues.

Swayed by the music: sampling bias towards musical preference distinguishes like from dislike decisions.

This study investigated the interaction between sampling behavior and preference formation underlying subjective decision making for like and dislike decisions. Two-alternative forced-choice tasks were used with closely-matched musical excerpts and the participants were free to listen and re-listen, i.e. to sample and resample each excerpt, until they reached a decision. We predicted that for decisions involving resampling, a sampling bias would be observed before the moment of conscious decision for the like decision only. The results indeed showed a gradually increasing sampling bias favouring the choice (73%) before the moment of overt response for like decisions. Such a bias was absent for dislike decisions. Furthermore, the participants reported stronger relative preferences for like decisions as compared to dislike decisions. This study demonstrated distinct differences in preference formation between like and dislike decisions, both in the implicit orienting/sampling processes prior to the conscious decision and in the subjective evaluation afterwards.

Team Flow Is a Unique Brain State Associated with Enhanced Information Integration and Interbrain Synchrony.

Team flow occurs when a group functions in a high task engagement to achieve a goal, commonly seen in performance and sports. Team flow can enable enhanced positive experiences, as compared with individual flow or regular socializing. However, the neural basis for this enhanced behavioral state remains unclear. Here, we identified neural correlates (NCs) of team flow in human participants using a music rhythm task with electroencephalogram hyperscanning. Experimental manipulations held the motor task constant while disrupting the corresponding hedonic music to interfere with the flow state or occluding the partner's positive feedback to impede team interaction. We validated these manipulations by using psychometric ratings and an objective measure for the depth of flow experience, which uses the auditory-evoked potential (AEP) of a task-irrelevant stimulus. Spectral power analysis at both the scalp sensors and anatomic source levels revealed higher ß-γ power specific to team flow in the left middle temporal cortex (L-MTC). Causal interaction analysis revealed that the L-MTC is downstream in information processing and receives information from areas encoding the flow or social states. The L-MTC significantly contributes to integrating information. Moreover, we found that team flow enhances global interbrain integrated information (II) and neural synchrony. We conclude that the NCs of team flow induce a distinct brain state. Our results suggest a neurocognitive mechanism to create this unique experience.