Implicit semantics gates visual awareness.

The current study asked whether impoverished peripheral vision led to perception immune from word-based semantic influences. We leveraged a peripheral sound-induced flash illusion. In each trial, two or three Mandarin characters were flashed quickly in the periphery with number-congruent or -incongruent beeps. We first successfully replicated the original illusions, showing auditory dominance. For example, when three characters were presented together with two beeps, the observer reported perceiving only two characters. Similarly, an additional beep induced an illusory visual percept. Crucially, when the three characters formed a meaningful word, the lack of a concurrent beep suppressed the awareness to a greater extent. A separate experiment replicated the effect on participants who were unable to recognize the words. When the reading was disrupted by reversing the presentation order, the effect disappeared. These findings demonstrate the capacity of our visual system to extract peripheral linguistic information without conscious word recognition.

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.

Detecting spontaneous deception in the brain.

Deception detection can be of great value during the juristic investigation. Although the neural signatures of deception have been widely documented, most prior studies were biased by difficulty levels. That is, deceptive behavior typically required more effort, making deception detection possibly effort detection. Furthermore, no study has examined the generalizability across instructed and spontaneous responses and across participants. To explore these issues, we used a dual-task paradigm, where the difficulty level was balanced between truth-telling and lying, and the instructed and spontaneous truth-telling and lying were collected independently. Using Multivoxel pattern analysis, we were able to decode truth-telling versus lying with a balanced difficulty level. Results showed that the angular gyrus (AG), inferior frontal gyrus (IFG), and postcentral gyrus could differentiate lying from truth-telling. Critically, linear classifiers trained to distinguish instructed truthful and deceptive responses could correctly differentiate spontaneous truthful and deceptive responses in AG and IFG with above-chance accuracy. In addition, with a leave-one-participant-out analysis, multivoxel neural patterns from AG could classify if the left-out participant was lying or not in a trial. These results indicate the commonality of neural responses subserved instructed and spontaneous deceptive behavior as well as the feasibility of cross-participant deception validation.

Subliminal temporal integration of linguistic information under discontinuous flash suppression.

Whether unconscious complex visual information integration occurs over time remains largely unknown and highly controversial. Previous studies have tended to use a combination of strong masking or suppression and a weak stimulus signal (e.g., low luminance), resulting in a low signal-to-noise ratio during unconscious stimulus presentation. To lengthen the stimulus exposure, we introduced intermittent presentation into interocular suppression. This discontinuous suppression allowed us to insert a word during each suppression period and deliver multiple words over time unconsciously. We found that, after participants received the subliminal context, they responded faster to a syntactically incongruent target word in a lexical decision task. We later replicated the finding in a separate experiment where participants exhibited chance performance on locating the subliminal context. These results confirmed that the sentential context was both subjectively and objectively subliminal. Critically, the effect disappeared when the context was disrupted by presenting only partial sentences or sentences with a reversed word order. These control experiments showed that the effect was not merely driven by word-word association but instead required integration over multiple words in the correct order. These findings support the possibility of unconscious high-level, complex information integration.

Cerebral neural correlates of differential melanopic photic stimulation in humans.

Photic stimulation of rods, cones and intrinsically photosensitive melanopsin-containing retinal ganglion cells (ipRGCs) mediates non-visual light responses, including entrainment of circadian rhythms and pupillary light reflex. Unlike visual responses to photic stimulation, the cerebral correlates of non-visual light responses in humans remains elusive. In this study, we used functional magnetic resonance imaging (fMRI) in 14 healthy young participants, to localize cerebral regions which are differentially activated by metameric light that gave rise to different levels of melanopic excitation. Mean blood oxygen-level dependent (BOLD) responses disclosed bilateral activation of the frontal eye fields during exposure to light geared towards melanopsin. Furthermore, multivariate pattern analyses showed distinct bilateral pattern activity in the inferior temporal gyri and the caudate nuclei. Taken together, our findings suggest that melanopsin-based photoreception activates a cerebral network including frontal regions, classically involved in attention and ocular motor responses.

Can a Word Sound Like a Shape Before You Have Seen It? Sound-Shape Mapping Prior to Conscious Awareness.

Nonarbitrary mappings between sound and shape (i.e., the bouba-kiki effect) have been shown across different cultures and early in development; however, the level of processing at which this effect arises remains unclear. Here we show that the mapping occurs prior to conscious awareness of the visual stimuli. Under continuous flash suppression, congruent stimuli (e.g., "kiki" inside an angular shape) broke through to conscious awareness faster than incongruent stimuli. This was true even when we trained people to pair unfamiliar letters with auditory word forms, a result showing that the effect was driven by the phonology, not the visual features, of the letters. Furthermore, visibility thresholds of the shapes decreased when they were preceded by a congruent auditory word form in a masking paradigm. Taken together, our results suggest that sound-shape mapping can occur automatically prior to conscious awareness of visual shapes, and that sensory congruence facilitates conscious awareness of a stimulus being present.

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.

Individual variability in neural representations of mind-wandering.

Mind-wandering is a frequent, daily mental activity, experienced in unique ways in each person. Yet neuroimaging evidence relating mind-wandering to brain activity, for example in the default mode network (DMN), has relied on population-rather than individual-based inferences due to limited within-individual sampling. Here, three densely-sampled individuals each reported hundreds of mind-wandering episodes while undergoing multi-session functional magnetic resonance imaging. We found reliable associations between mind-wandering and DMN activation when estimating brain networks within individuals using precision functional mapping. However, the timing of spontaneous DMN activity relative to subjective reports, and the networks beyond DMN that were activated and deactivated during mind-wandering, were distinct across individuals. Connectome-based predictive modeling further revealed idiosyncratic, whole-brain functional connectivity patterns that consistently predicted mind-wandering within individuals but did not fully generalize across individuals. Predictive models of mind-wandering and attention that were derived from larger-scale neuroimaging datasets largely failed when applied to densely-sampled individuals, further highlighting the need for personalized models. Our work offers novel evidence for both conserved and variable neural representations of self-reported mind-wandering in different individuals. The previously-unrecognized inter-individual variations reported here underscore the broader scientific value and potential clinical utility of idiographic approaches to brain-experience associations.

Individual variability in neural representations of mind-wandering.

Mind-wandering is a frequent, daily mental activity, experienced in unique ways in each person. Yet neuroimaging evidence relating mind-wandering to brain activity, for example in the default mode network (DMN), has relied on population- rather than individual-based inferences owing to limited within-person sampling. Here, three densely sampled individuals each reported hundreds of mind-wandering episodes while undergoing multi-session functional magnetic resonance imaging. We found reliable associations between mind-wandering and DMN activation when estimating brain networks within individuals using precision functional mapping. However, the timing of spontaneous DMN activity relative to subjective reports, and the networks beyond DMN that were activated and deactivated during mind-wandering, were distinct across individuals. Connectome-based predictive modeling further revealed idiosyncratic, whole-brain functional connectivity patterns that consistently predicted mind-wandering within individuals but did not fully generalize across individuals. Predictive models of mind-wandering and attention that were derived from larger-scale neuroimaging datasets largely failed when applied to densely sampled individuals, further highlighting the need for personalized models. Our work offers novel evidence for both conserved and variable neural representations of self-reported mind-wandering in different individuals. The previously unrecognized interindividual variations reported here underscore the broader scientific value and potential clinical utility of idiographic approaches to brain-experience associations.

While everyone experiences that their mind "wanders" throughout daily life, the content and form of inner experience is different in different people. In this study, we found that brain activity representing mind-wandering is different in each person, reflecting unique mental experiences. While people consistently engaged the brain's default mode network (DMN) during mind-wandering, there were inconsistencies in the way that the DMN was engaged and in the other networks throughout the brain that were engaged. Our study highlights that personalized approaches, which require that individuals are sampled more densely than is common in current practice, enable accurate insights into relationships between brain activity and inner experience.

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.

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.

Alpha desynchronization during Stroop test unmasks cognitively healthy individuals with abnormal CSF Amyloid/Tau.

Synaptic dysfunctions precede cognitive decline in Alzheimer's disease by decades, affect executive functions, and can be detected by quantitative electroencephalography (qEEG). We used quantitative electroencephalography combined with Stroop testing to identify changes of inhibitory controls in cognitively healthy individuals with an abnormal versus normal ratio of cerebrospinal fluid (CSF) amyloid/total-tau. We studied two groups of participants (60-94 years) with either normal (CH-NAT or controls, n = 20) or abnormal (CH-PAT, n = 21) CSF amyloid/tau ratio. We compared: alpha event-related desynchronization (ERD), alpha spectral entropy (SE), and their relationships with estimated cognitive reserve. CH-PATs had more negative occipital alpha ERD, and higher frontal and occipital alpha SE during low load congruent trials, indicating hyperactivity. CH-PATs demonstrated fewer frontal SE changes with higher load, incongruent Stroop testing. Correlations of alpha ERD with estimated cognitive reserve were significant in CH-PATs but not in CH-NATs. These results suggested compensatory hyperactivity in CH-PATs compared to CH-NATs. We did not find differences in alpha ERD comparisons with individual CSF amyloid(A), p-tau(T), total-tau(N) biomarkers.

Task switching reveals abnormal brain-heart electrophysiological signatures in cognitively healthy individuals with abnormal CSF amyloid/tau, a pilot study.

Electroencephalographic (EEG) alpha oscillations have been related to heart rate variability (HRV) and both change in Alzheimer's disease (AD). We explored if task switching reveals altered alpha power and HRV in cognitively healthy individuals with AD pathology in cerebrospinal fluid (CSF) and whether HRV improves the AD pathology classification by alpha power alone. We compared low and high alpha event-related desynchronization (ERD) and HRV parameters during task switch testing between two groups of cognitively healthy participants classified by CSF amyloid/tau ratio: normal (CH-NAT, n = 19) or pathological (CH-PAT, n = 27). For the task switching paradigm, participants were required to name the color or word for each colored word stimulus, with two sequential stimuli per trial. Trials include color (cC) or word (wW) repeats with low load repeating, and word (cW) or color switch (wC) for high load switching. HRV was assessed for RR interval, standard deviation of RR-intervals (SDNN) and root mean squared successive differences (RMSSD) in time domain, and low frequency (LF), high frequency (HF), and LF/HF ratio in frequency domain. Results showed that CH-PATs compared to CH-NATs presented: 1) increased (less negative) low alpha ERD during low load repeat trials and lower word switch cost (low alpha: p = 0.008, Cohen's d = -0.83, 95% confidence interval -1.44 to -0.22, and high alpha: p = 0.019, Cohen's d = -0.73, 95% confidence interval -1.34 to -0.13); 2) decreasing HRV from rest to task, suggesting hyper-activated sympatho-vagal responses. 3) CH-PATs classification by alpha ERD was improved by supplementing HRV signatures, supporting a potentially compromised brain-heart interoceptive regulation in CH-PATs. Further experiments are needed to validate these findings for clinical significance.

Task-induced attention load guides and gates unconscious semantic interference.

The tight relationship between attention and conscious perception has been extensively researched in the past decades. However, whether attentional modulation extended to unconscious processes remained largely unknown, particularly when it came to abstract and high-level processing. Here we use a double Stroop paradigm to demonstrate that attention load gates unconscious semantic processing. We find that word and color incongruencies between a subliminal prime and a supraliminal target cause slower responses to non-Stroop target words-but only if the task is to name the target word (low-load task), and not if the task is to name the target's color (high-load task). The task load hypothesis is confirmed by showing that the word-induced incongruence effect can be detected in the color-naming task, but only in the late, practiced trials. We further replicate this task-induced attentional modulation phenomenon in separate experiments with colorless words (word-only) and words with semantic relationship but no orthographic similarities (semantics-only).

Unconscious processing of facial attractiveness: invisible attractive faces orient visual attention.

Past research has proven human's extraordinary ability to extract information from a face in the blink of an eye, including its emotion, gaze direction, and attractiveness. However, it remains elusive whether facial attractiveness can be processed and influences our behaviors in the complete absence of conscious awareness. Here we demonstrate unconscious processing of facial attractiveness with three distinct approaches. In Experiment 1, the time taken for faces to break interocular suppression was measured. The results showed that attractive faces enjoyed the privilege of breaking suppression and reaching consciousness earlier. In Experiment 2, we further showed that attractive faces had lower visibility thresholds, again suggesting that facial attractiveness could be processed more easily to reach consciousness. Crucially, in Experiment 3, a significant decrease of accuracy on an orientation discrimination task subsequent to an invisible attractive face showed that attractive faces, albeit suppressed and invisible, still exerted an effect by orienting attention. Taken together, for the first time, we show that facial attractiveness can be processed in the complete absence of consciousness, and an unconscious attractive face is still capable of directing our attention.

Syntactic processing in the absence of awareness and semantics.

The classical view that multistep rule-based operations require consciousness has recently been challenged by findings that both multiword semantic processing and multistep arithmetic equations can be processed unconsciously. It remains unclear, however, whether pure rule-based cognitive processes can occur unconsciously in the absence of semantics. Here, after presenting 2 words consciously, we suppressed the third with continuous flash suppression. First, we showed that the third word in the subject-verb-verb format (syntactically incongruent) broke suppression significantly faster than the third word in the subject-verb-object format (syntactically congruent). Crucially, the same effect was observed even with sentences composed of pseudowords (pseudo subject-verb-adjective vs. pseudo subject-verb-object) without any semantic information. This is the first study to show that syntactic congruency can be processed unconsciously in the complete absence of semantics. Our findings illustrate how abstract rule-based processing (e.g., syntactic categories) can occur in the absence of visual awareness, even when deprived of semantics.

The role of inferior frontal gyrus in processing Chinese classifiers.

The Chinese classifier system classifies nouns and builds a relation between classifiers and their corresponding nouns. This functional magnetic resonance imaging (fMRI) study examined brain activation of Chinese classifiers during reading comprehension. Thirty-four participants read and performed semantic congruency judgments on congruent, inside-classifier (IC) violated, and outside-classifier (OC) violated sentences. The IC and OC violations were created by changing the correct classifier to an inappropriate classifier and a non-classifier, respectively. The comparison of the IC violation vs. the congruent condition produced greater activation in the mid-ventral region (BA 45) of the left inferior frontal gyrus (IFG), suggesting an increased demand on semantic processing. Contrasting different subtypes of IC violation produced greater activation in the right IFG (BAs 45 and 47), indicating that processing mass/count classifiers involves distinct brain activations. The OC violation produced greater activation in the left IFG (BAs 45 and 44), suggesting both semantic and syntactic processing. These results indicate that different parts of the IFG contribute to syntactic and semantic processing of classifier phrases in reading Chinese for comprehension.