Different kinds of simulation during literary reading: Insights from a combined fMRI and eye-tracking study.

Mental simulation is an important aspect of narrative reading. In a previous study, we found that gaze durations are differentially impacted by different kinds of mental simulation. Motor simulation, perceptual simulation, and mentalizing as elicited by literary short stories influenced eye movements in distinguishable ways (Mak & Willems, 2019). In the current study, we investigated the existence of a common neural locus for these different kinds of simulation. We additionally investigated whether individual differences during reading, as indexed by the eye movements, are reflected in domain-specific activations in the brain. We found a variety of brain areas activated by simulation-eliciting content, both modality-specific brain areas and a general simulation area. Individual variation in percent signal change in activated areas was related to measures of story appreciation as well as personal characteristics (i.e., transportability, perspective taking). Taken together, these findings suggest that mental simulation is supported by both domain-specific processes grounded in previous experiences, and by the neural mechanisms that underlie higher-order language processing (e.g., situation model building, event indexing, integration).

Different routes to liking: how readers arrive at narrative evaluations.

When two people read the same story, they might both end up liking it very much. However, this does not necessarily mean that their reasons for liking it were identical. We therefore ask what factors contribute to "liking" a story, and-most importantly-how people vary in this respect. We found that readers like stories because they find them interesting, amusing, suspenseful and/or beautiful. However, the degree to which these components of appreciation were related to how much readers liked stories differed between individuals. Interestingly, the individual slopes of the relationships between many of the components and liking were (positively or negatively) correlated. This indicated, for instance, that individuals displaying a relatively strong relationship between interest and liking, generally display a relatively weak relationship between sadness and liking. The individual differences in the strengths of the relationships between the components and liking were not related to individual differences in expertize, a characteristic strongly associated with aesthetic appreciation of visual art. Our work illustrates that it is important to take into consideration the fact that individuals differ in how they arrive at their evaluation of literary stories, and that it is possible to quantify these differences in empirical experiments. Our work suggests that future research should be careful about "overfitting" theories of aesthetic appreciation to an "idealized reader," but rather take into consideration variations across individuals in the reason for liking a particular story.

Correction: Protocol of the Healthy Brain Study: An accessible resource for understanding the human brain and how it dynamically and individually operates in its bio-social context.

[This corrects the article DOI: 10.1371/journal.pone.0260952.].

Mapping dopaminergic projections in the human brain with resting-state fMRI.

The striatum receives dense dopaminergic projections, making it a key region of the dopaminergic system. Its dysfunction has been implicated in various conditions including Parkinson's disease (PD) and substance use disorder. However, the investigation of dopamine-specific functioning in humans is problematic as current MRI approaches are unable to differentiate between dopaminergic and other projections. Here, we demonstrate that 'connectopic mapping' - a novel approach for characterizing fine-grained, overlapping modes of functional connectivity - can be used to map dopaminergic projections in striatum. We applied connectopic mapping to resting-state functional MRI data of the Human Connectome Project (population cohort; N = 839) and selected the second-order striatal connectivity mode for further analyses. We first validated its specificity to dopaminergic projections by demonstrating a high spatial correlation (r = 0.884) with dopamine transporter availability - a marker of dopaminergic projections - derived from DaT SPECT scans of 209 healthy controls. Next, we obtained the subject-specific second-order modes from 20 controls and 39 PD patients scanned under placebo and under dopamine replacement therapy (L-DOPA), and show that our proposed dopaminergic marker tracks PD diagnosis, symptom severity, and sensitivity to L-DOPA. Finally, across 30 daily alcohol users and 38 daily smokers, we establish strong associations with self-reported alcohol and nicotine use. Our findings provide evidence that the second-order mode of functional connectivity in striatum maps onto dopaminergic projections, tracks inter-individual differences in PD symptom severity and L-DOPA sensitivity, and exhibits strong associations with levels of nicotine and alcohol use, thereby offering a new biomarker for dopamine-related (dys)function in the human brain.

Protocol of the Healthy Brain Study: An accessible resource for understanding the human brain and how it dynamically and individually operates in its bio-social context.

The endeavor to understand the human brain has seen more progress in the last few decades than in the previous two millennia. Still, our understanding of how the human brain relates to behavior in the real world and how this link is modulated by biological, social, and environmental factors is limited. To address this, we designed the Healthy Brain Study (HBS), an interdisciplinary, longitudinal, cohort study based on multidimensional, dynamic assessments in both the laboratory and the real world. Here, we describe the rationale and design of the currently ongoing HBS. The HBS is examining a population-based sample of 1,000 healthy participants (age 30-39) who are thoroughly studied across an entire year. Data are collected through cognitive, affective, behavioral, and physiological testing, neuroimaging, bio-sampling, questionnaires, ecological momentary assessment, and real-world assessments using wearable devices. These data will become an accessible resource for the scientific community enabling the next step in understanding the human brain and how it dynamically and individually operates in its bio-social context. An access procedure to the collected data and bio-samples is in place and published on https://www.healthybrainstudy.nl/en/data-and-methods/access. Trail registration: https://www.trialregister.nl/trial/7955.

The Impact of Error-Consequence Severity on Cue Processing in Importance-Biased Prospective Memory.

Prospective memory (PM) enables people to remember to complete important tasks in the future. Failing to do so can result in consequences of varying severity. Here, we investigated how PM error-consequence severity impacts the neural processing of relevant cues for triggering PM and the ramification of that processing on the associated prospective task performance. Participants role-played a cafeteria worker serving lunches to fictitious students and had to remember to deliver an alternative lunch to students (as PM cues) who would otherwise experience a moderate or severe aversive reaction. Scalp-recorded, event-related potential (ERP) measures showed that the early-latency frontal positivity, reflecting the perception-based neural responses to previously learned stimuli, did not differ between the severe versus moderate PM cues. In contrast, the longer-latency parietal positivity, thought to reflect full PM cue recognition and post-retrieval processes, was elicited earlier by the severe than the moderate PM cues. This faster instantiation of the parietal positivity to the severe-consequence PM cues was then followed by faster and more accurate behavioral responses. These findings indicate how the relative importance of a PM can be neurally instantiated in the form of enhanced and faster PM-cue recognition and processing and culminate into better PM.

Functional co-activation of the default mode network in APOE ε4-carriers: A replication study.

Structural and functional alterations of the brain in persons genetically at-risk for Alzheimer's disease (AD) are crucial in unravelling AD development. Filippini et al. found that the default mode network (DMN) is already affected in young APOE ε4-carriers, with increased co-activation of the DMN during rest and increased hippocampal task activation. We aimed to replicate the early findings of Filippini et al, using the APOE gene, still the principal AD risk gene, and extended this with a polygenic risk score (PRS) analysis for AD, using the Human Connectome Project dataset (HCP). We included participants from the HCP S1200 dataset (age range: 22-36 years). We studied morphometric features, functional DMN co-activation and functional task activation of recollection performance. Permutation Analysis of Linear Models (PALM) was used to test for group differences between APOE ε4-carriers and non-carriers, and to test the association with PRS. PALM controls for biases induced by the family structure of the HCP sample. Results were family-wise error rate corrected at p < 0.05. Our primary analysis did not replicate the early findings of Filippini et al. (2009). However, compared with non-carriers, APOE ε4-carriers showed increased functional activation during the encoding of subsequently recollected items in areas related to facial recognition (p<0.05, t>756.11). This increased functional activation was also positively associated with PRS (APOE variants included) (p<0.05, t>647.55). Our results are supportive for none to limited genetic effects on brain structure and function in young adults. Taking the methodological considerations of replication studies into account, the true effect of APOE ε4-carriership is likely smaller than indicated in the Filippini paper. However, it still holds that we may not yet be able to detect already present measurable effects decades before a clinical expression of AD. Since the mechanistic pathway of AD is likely to encompass many different factors, further research should be focused on the interactions of genetic risk, biomarkers, aging and lifestyle factors over the life course. Sensitive functional neuroimaging as used here may help disentangling these complex interactions.

Where the eyes wander: The relationship between mind wandering and fixation allocation to visually salient and semantically informative static scene content.

Vision is crucial for many everyday activities, but the mind is not always focused on what the eyes see. Mind wandering occurs frequently and is associated with attenuated visual and cognitive processing of external information. Corresponding changes in gaze behavior-namely, fewer, longer, and more dispersed fixations-suggest a shift in how the visual system samples external information. Using three computational models of visual salience and two innovative approaches for measuring semantic informativeness, the current work assessed whether these changes reflect how the visual system prioritizes visually salient and semantically informative scene content, two major determinants in most theoretical frameworks and computational models of gaze control. Findings showed that, in a static scene viewing task, fixations were allocated to scene content that was more visually salient 10 seconds prior to probe-caught, self-reported mind wandering compared to self-reported attentive viewing. The relationship between mind wandering and semantic content was more equivocal, with weaker evidence that fixations are more likely to fall on locally informative scene regions. This indicates that the visual system is still able to discriminate visually salient and semantically informative scene content during mind wandering and may fixate on such information more frequently than during attentive viewing. Theoretical implications are discussed in light of these findings.

Mind wandering as data augmentation: How mental travel supports abstraction.

Gilead et al. state that abstraction supports mental travel, and that mental travel critically relies on abstraction. I propose an important addition to this theoretical framework, namely that mental travel might also support abstraction. Specifically, I argue that spontaneous mental travel (mind wandering), much like data augmentation in machine learning, provides variability in mental content and context necessary for abstraction.

The eye-mind wandering link: Identifying gaze indices of mind wandering across tasks.

During mind wandering, visual processing of external information is attenuated. Accordingly, mind wandering is associated with changes in gaze behaviors, albeit findings are inconsistent in the literature. This heterogeneity obfuscates a complete view of the moment-to-moment processing priorities of the visual system during mind wandering. We hypothesize that this observed heterogeneity is an effect of idiosyncrasy across tasks with varying spatial allocation demands, visual processing demands, and discourse processing demands and reflects a strategic, compensatory shift in how the visual system operates during mind wandering. We recorded eye movements and mind wandering (via thought-probes) as 132 college-aged adults completed a battery of 7 short (6 min) tasks with different visual demands. We found that for tasks requiring extensive sampling of the visual field, there were fewer fixations, and, depending on the specific task, fixations were longer and/or more dispersed. This suggests that visual sampling is sparser and potentially slower and more dispersed to compensate for the decreased sampling rate during mind wandering. For tasks that demand centrally focused gaze, mind wandering was accompanied by more exploratory eye movements, such as shorter and more dispersed fixations as well as larger saccades. Gaze behaviors were not reliably associated with mind wandering during a film comprehension task. These findings provide insight into how the visual system prioritizes external information when attention is focused inward and indicates the importance of task demands when assessing the relationship between eye movements, visual processing, and mind wandering. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Word skipping as an indicator of individual reading style during literary reading.

Decades of research have established that the content of language (e.g. lexical characteristics of words) predicts eye movements during reading. Here we investigate whether there exist individual differences in 'stable' eye movement patterns during narrative reading. We computed Euclidean distances from correlations between gaze durations time courses (word level) across 102 participants who each read three literary narratives in Dutch. The resulting distance matrices were compared between narratives using a Mantel test. The results show that correlations between the scaling matrices of different narratives are relatively weak (r ≤ .11) when missing data points are ignored. However, when including these data points as zero durations (i.e. skipped words), we found significant correlations between stories (r > .51). Word skipping was significantly positively associated with print exposure but not with self-rated attention and story-world absorption, suggesting that more experienced readers are more likely to skip words, and do so in a comparable fashion. We interpret this finding as suggesting that word skipping might be a stable individual eye movement pattern.

The functional organisation of the hippocampus along its long axis is gradual and predicts recollection.

Understanding the functional organisation of the hippocampus is crucial for understanding its role in cognition and disorders in which it is implicated. Different views have been proposed of how function is distributed along its long axis: one view suggests segregation, whereas the alternative view postulates a more gradual organisation. Here, we applied a novel 'connectopic mapping' data-analysis approach to the resting-state fMRI data of participants of the Human Connectome Project, and demonstrate that the functional organisation of the hippocampal longitudinal axis is gradual rather than segregated into parcels. In addition, we show that inter-individual variations in this gradual organisation predict variations in recollection memory better than a characterisation based on functional parcellation. These results present an important step forward in understanding the functional organisation of the human hippocampus and have important implications for translating between rodent and human research.

How the stimulus influences mind wandering in semantically rich task contexts.

What do we think about when we mind wander and where do these thoughts come from? We tested the idea that semantically rich stimuli yield patterns of mind wandering that are closely coupled with the stimuli compared to being more internally triggered. We analyzed the content of 949 self-reported zone outs (1218 thoughts) and 519 of their triggers from 88 participants who read an instructional text and watched a film for 20 min each. We found that mind wandering associated with memory retrieval was more frequent than prospection and introspection across both stimuli. Over 70% of autobiographical and semantic memory retrievals were triggered by the content of the stimuli, compared to around 30% for prospective and introspective thoughts. Further, latent semantic analysis revealed that semantic and unspecific memories were more "semantically" similar to their triggers than prospective and introspective thoughts, suggesting that they arise from spontaneous associations with the stimulus. These findings suggest a re-evaluation of how internal concerns and the external world give rise to mind wandering and emphasize the importance of studying mind wandering in semantically rich contexts akin to much of the real world.

Gaze-based signatures of mind wandering during real-world scene processing.

Physiological limitations on the visual system require gaze to move from location to location to extract the most relevant information within a scene. Therefore, gaze provides a real-time index of the information-processing priorities of the visual system. We investigated gaze allocation during mind wandering (MW), a state where cognitive priorities shift from processing task-relevant external stimuli (i.e., the visual world) to task-irrelevant internal thoughts. In both a main study and a replication, we recorded the eye movements of college-aged adults who studied images of urban scenes and responded to pseudorandom thought probes on whether they were mind wandering or attentively viewing at the time of the probe. Probe-caught MW was associated with fewer and longer fixations, greater fixation dispersion, and more frequent eyeblinks (only observed in the main study) relative to periods of attentive scene viewing. These findings demonstrate that gaze indices typically considered to represent greater engagement with scene processing (e.g., longer fixations) can also indicate MW. In this way, the current work exhibits a need for empirical investigations and computational models of gaze control to account for MW for a more accurate representation of the moment-to-moment information-processing priorities of the visual system. (PsycINFO Database Record

Driven to distraction: A lack of change gives rise to mind wandering.

How does the dynamic structure of the external world direct attention? We examined the relationship between event structure and attention to test the hypothesis that narrative shifts (both theoretical and perceived) negatively predict attentional lapses. Self-caught instances of mind wandering were collected while 108 participants watched a 32.5 min film called The Red Balloon. We used theoretical codings of situational change and human perceptions of event boundaries to predict mind wandering in 5-s intervals. Our findings suggest a temporal alignment between the structural dynamics of the film and mind wandering reports. Specifically, the number of situational changes and likelihood of perceiving event boundaries in the prior 0-15 s interval negatively predicted mind wandering net of low-level audiovisual features. Thus, mind wandering is less likely to occur when there is more event change, suggesting that narrative shifts keep attention from drifting inwards.

An automated behavioral measure of mind wandering during computerized reading.

Mind wandering is a ubiquitous phenomenon in which attention shifts from task-related to task-unrelated thoughts. The last decade has witnessed an explosion of interest in mind wandering, but research has been stymied by a lack of objective measures, leading to a near-exclusive reliance on self-reports. We addressed this issue by developing an eye-gaze-based, machine-learned model of mind wandering during computerized reading. Data were collected in a study in which 132 participants reported self-caught mind wandering while reading excerpts from a book on a computer screen. A remote Tobii TX300 or T60 eyetracker recorded their gaze during reading. The data were used to train supervised classification models to discriminate between mind wandering and normal reading in a manner that would generalize to new participants. We found that at the point of maximal agreement between the model-based and self-reported mind-wandering means (smallest difference between the group-level means: M model = .310, M self = .319), the participant-level mind-wandering proportional distributions were similar and were significantly correlated (r = .400). The model-based estimates were internally consistent (r = .751) and predicted text comprehension more strongly than did self-reported mind wandering (r model = -.374, r self = -.208). Our results also indicate that a robust strategy of probabilistically predicting mind wandering in cases with poor or missing gaze data led to improved performance on all metrics, as compared to simply discarding these data. Our findings demonstrate that an automated objective measure might be available for laboratory studies of mind wandering during reading, providing an appealing alternative or complement to self-reports.

Effects of learned episodic event structure on prospective duration judgments.

The field of psychology of time has typically distinguished between prospective timing and retrospective duration estimation: in prospective timing, participants attend to and encode time, whereas in retrospective estimation, estimates are based on the memory of what happened. Prior research on prospective timing has primarily focused on attentional mechanisms to explain timing behavior, but it remains unclear the extent to which memory processes may also play a role. The present studies investigate this issue, and specifically, the role of newly learned encoded event structure. Two structural properties of dynamic event sequences were examined, which are known to modulate retrospective duration estimates: the perceived number of segments and the similarity between them. We found that when duration and episodic event content are both attended to and encoded, more segments and less similarity between them led to longer attributed durations, despite clock duration remaining constant. In contrast, when only duration is attended to, only the number of segments influenced estimated durations. These findings indicate that incidentally or intentionally encoded episodic event structure modulates prospective duration judgments. Based on these and previous findings, implications for the role of memory mechanisms on prospective paradigms are discussed. (PsycINFO Database Record

The effect of disfluency on mind wandering during text comprehension.

When reading, we frequently find ourselves thinking about something other than the text. These attentional lapses, known as mind wandering (MW), are negatively correlated with text comprehension. Previous studies have shown that more syntactically and semantically difficult texts elicit more MW, because textual difficulty impedes the construction of a mental model of the text, which makes it more difficult to suppress off-task thoughts. But is it possible to reduce MW without altering the content of the text itself? We hypothesized that reading a perceptually disfluent text might require more attentional resources, even if the content remained the same, leaving fewer resources available for MW. To test this idea, we manipulated the typefaces (fluent [Arial] or disfluent [ ]) of two instructional texts on scientific research methods (each about 1,490 words long), and found that MW was less frequent when participants read the disfluent text. There were no comprehension differences between the fluent and disfluent groups. However, we did find an indirect effect of disfluency on comprehension through MW, suggesting that disfluency influences comprehension by enhancing attention. These findings provide insights into how processing difficulty and attention interact during reading comprehension.