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Reducing contributions from non-neuronal sources is a crucial step in functional magnetic resonance imaging (fMRI) analyses. Many viable strategies for denoising fMRI are used in the literature, and practitioners rely on denoising benchmarks for guidance in the selection of an appropriate choice for their study. However, fMRI denoising software is an ever-evolving field, and the benchmarks can quickly become obsolete as the techniques or implementations change. In this work, we present a fully reproducible denoising benchmark featuring a range of denoising strategies and evaluation metrics, built primarily on the fMRIPrep and Nilearn software packages. We apply this reproducible benchmark to investigate the robustness of the conclusions across two different datasets and two versions of fMRIPrep. The majority of benchmark results were consistent with prior literature. Scrubbing, a technique which excludes time points with excessive motion, combined with global signal regression, is generally effective at noise removal. Scrubbing however disrupts the continuous sampling of brain images and is incompatible with some statistical analyses, e.g. auto-regressive modeling. In this case, a simple strategy using motion parameters, average activity in select brain compartments, and global signal regression should be preferred. Importantly, we found that certain denoising strategies behave inconsistently across datasets and/or versions of fMRIPrep, or had a different behavior than in previously published benchmarks, especially ICA-AROMA. These results demonstrate that a reproducible denoising benchmark can effectively assess the robustness of conclusions across multiple datasets and software versions. Technologies such as BIDS-App, the Jupyter Book and Neurolibre provided the infrastructure to publish the metadata and report figures. Readers can reproduce the report figures beyond the ones reported in the published manuscript. With the denoising benchmark, we hope to provide useful guidelines for the community, and that our software infrastructure will facilitate continued development as the state-of-the-art advances.

Insofar as the autistic-like phenotype presents in the general population, it consists of partially dissociable traits, such as social and sensory issues. Here, we investigate individual differences in cortical organisation related to autistic-like traits. Connectome gradient decomposition based on resting state fMRI data reliably reveals a principal gradient spanning from unimodal to transmodal regions, reflecting the transition from perception to abstract cognition. In our non-clinical sample, this gradient's expansion, indicating less integration between visual and default mode networks, correlates with subjective sensory sensitivity (measured using the Glasgow Sensory Questionnaire, GSQ), but not other autistic-like traits (measured using the Autism Spectrum Quotient, AQ). This novel brain-based correlate of the GSQ demonstrates sensory issues can be disentangled from the wider autistic-like phenotype.

Background Auditory verbal hallucinations (AVH) commonly occur in the context of borderline personality disorder (BPD) yet remain poorly understood. AVH are often perceived by patients with BPD as originating from inside the head and hence viewed clinically as “pseudohallucinations,” but they nevertheless have a detrimental impact on well-being. Methods The current study characterized perceptual, subjective, and neural expressions of AVH by using an auditory detection task, experience sampling and questionnaires, and functional neuroimaging, respectively. Results Perceptually, reported AVH correlated with a bias for reporting the presence of a voice in white noise. Subjectively, questionnaire measures indicated that AVH were significantly distressing and persecutory. In addition, AVH intensity, but not perceived origin (i.e., inside vs outside the head), was associated with greater concurrent anxiety. Neurally, fMRI of BPD participants demonstrated that, relative to imagining or listening to voices, periods of reported AVH induced greater blood oxygenation level–dependent activity in anterior cingulate and bilateral temporal cortices (regional substrates for language processing). AVH symptom severity was associated with weaker functional connectivity between anterior cingulate and bilateral insular cortices. Conclusion In summary, our results indicate that AVH in participants with BPD are (1) underpinned by aberrant perceptual-cognitive mechanisms for signal detection, (2) experienced subjectively as persecutory and distressing, and (3) associated with distinct patterns of neural activity that inform proximal mechanistic understanding. Our findings are like analogous observations in patients with schizophrenia and validate the clinical significance of the AVH experience in BPD, often dismissed as “pseudohallucinations.” These highlight a need to reconsider this experience as a treatment priority.

Objective To explore the protective effect of Qihuo decoction (QHD) on the livers of sleep-deprived rats housed in the humid and hot environment, and to characterize ultrastructural changes of livers in these rats. Methods Forty male SD rats were randomized into 4 groups (10 each group): blank control group (BC group), humid and hot control group (HC group), humid and hot with sleep deprivation group (HS group), and humid and hot with sleep-deprived treated with QHD group (HSQ group). Rats in BC group were housed in the normal temperature and humidity chamber; while rats in HC group, HS group, and HSQ group were housed in the simulated hot and humid chamber. Rats in BC group, HC group, and HS group were given distilled water to gavage for 15 days, and rats in HSQ group were given QHD to gavage for 15 days. Then rats in HS group and HSQ group were sleep deprived for 72 hours. The serum concentrations of alanine aminotransferase (ALT) and serum aspartate aminotransferase (AST) were detected. The liver tissues contents of hepatic glycogen, malondialdehyde (MDA) and reduced glutathione (GSH) were detected. The expression of β-galactosidase (β-gal) in each group was also detected. Lastly, the rat hepatocytes ultrastructure was observed by a transmission electron microscope. Results In BC group, HC group, HS group, and HSQ group, the serum AST levels were (107.7±8.4)U/L, (126.9±10.4)U/L, (176.9±13.1)U/L, and (140.6±9.6)U/L; the ALT levels were (37.9±3.1)U/L, (45.5±6.2)U/L, (86.5±10.4)U/L, and (64.9±8.7)U/L. Statistical analysis among groups were significantly different (P

Summary A core goal in cognitive neuroscience is identifying the physical substrates of the patterns of thought that occupy our daily lives. Contemporary views suggest that the landscape of ongoing experience is heterogeneous and can be influenced by features of both the person and the context. This perspective piece considers recent work that explicitly accounts for both the heterogeneity of the experience and context dependence of patterns of ongoing thought. These studies reveal that systems linked to attention and control are important for organizing experience in response to changing environmental demands. These studies also establish a role of the default mode network beyond task-negative or purely episodic content, for example, implicating it in the level of vivid detail in experience in both task contexts and in spontaneous self-generated experiential states. Together, this work demonstrates that the landscape of ongoing thought is reflected in the activity of multiple neural systems, and it is important to distinguish between processes contributing to how the experience unfolds from those linked to how these experiences are regulated., Cognitive Neuroscience; Psychology; Techniques in Neuroscience

There is a long history of, and renewed interest in, cardiac timing effects on behaviour and cognition. Cardiac timing effects may be identified by expressing events as a function of their location in the cardiac cycle, and applying circular (i.e. directional) statistics to test cardiac time-behaviour associations. Typically this approach 'stretches' all points in the cardiac cycle equally, but this is not necessarily physiologically valid. Moreover, many tests impose distributional assumptions that are not met by such data. We present a set of statistical techniques robust to this, instantiated within our new Cardiac Timing Toolbox (CaTT) for MATLAB: A physiologically-motivated method of wrapping behaviour to the cardiac cycle; and a set of non-parametric statistical tests that control for common confounds and distributional characteristics of these data. Using a reanalysis of previously published data, we guide readers through analyses using CaTT, aiding researchers in identifying physiologically plausible associations between heart-timing and cognition.

1ABSTRACTFeatures of ongoing experience are common across individuals and cultures. However, certain people express specific patterns to a greater extent than others. The current psychological theory assumes that individual differences in thought patterns occur because different types of experience depend on differences in associated neurocognitive mechanisms. Consequently, individual variation in the underlying neurocognitive architecture is hypothesised to determine the ease with which certain thought patterns are generated or maintained. Our study (N=178) tested this hypothesis using multivariate pattern analysis to infer shared variance among measures of cognitive function and neural organisation and examined whether these hidden structures explain reports of the patterns of on-going thoughts people experienced in the lab. We found that relatively better performance on tasks requiring primarily semantic knowledge, rather than executive control, was linked to a neural functional organisation that was associated, via meta-analysis, with task labels related to semantic associations (sentence processing, reading and verbal semantics). Variability of this functional mode predicted significant individual variation in the types of thoughts that individuals experienced in the laboratory: Neurocognitive patterns linked to better performance at tasks that required guidance from semantic representation, rather than those dependent on executive control, were associated with patterns of thought characterised by greater subjective detail and a focus on time periods other than the here and now. These relationships were consistent across different days and did not vary with task condition, indicating they are relatively stable features of an individual’s cognitive profile. Together these data confirm that individual variation in aspects of ongoing experience can be inferred from hidden neurocognitive architecture and demonstrate that performance trade-offs between executive control and long term semantic knowledge are linked to a person’s tendency to imagine situations that transcend the here and now.

Cognition is dynamic and involves both the maintenance of and transitions between neurocognitive states. While recent research has identified some of the neural systems involved in sustaining task states, it is less well understood how intrinsic influences on cognition emerge over time. The current study uses fMRI and Multi-Dimensional Experience Sampling (MDES) to chart how cognition changes over time from moments in time when external attention was established. We found that the passage of time was associated with brain regions associated with external attention decreasing in activity over time. Comparing this pattern of activity to defined functional hierarchies of brain organization, we found that it could be best understood as movement away from systems involved in task performance. Moments where the participants described their thoughts as off-task showed a significant similarity to the task-negative end of the same hierarchy. Finally, the greater the similarity of a participant's neural dynamics to this hierarchy the faster their rate of increasing off-task thought over time. These findings suggest topographical changes in neural processing that emerge over time and those seen during off-task thought can both be understood as a common shift away from neural motifs seen during complex task performance.

Cognition is dynamic and involves both the maintenance of and transitions between neurocognitive states. While recent research has identified some of the neural systems involved in sustaining task states, it is less well understood how intrinsic influences on cognition emerge over time. The current study uses fMRI and Multi-Dimensional Experience Sampling (MDES) to chart how cognition changes over time from moments in time when external attention was established. We found that the passage of time was associated with brain regions associated with external attention decreasing in activity over time. Comparing this pattern of activity to defined functional hierarchies of brain organization, we found that it could be best understood as movement away from systems involved in task performance. Moments where the participants described their thoughts as off-task showed a significant similarity to the task-negative end of the same hierarchy. Finally, the greater the similarity of a participant’s neural dynamics to this hierarchy the faster their rate of increasing off-task thought over time. These findings suggest topographical changes in neural processing that emerge over time and those seen during off-task thought can both be understood as a common shift away from neural motifs seen during complex task performance.

Conscious awareness of the world fluctuates, either through variation in the quality with which we perceive the environment, or, when attention switches to information generated using imagination rather than the external environment. Our study combined individual differences in experience sampling, psychophysical reports of perception, and neuroimaging descriptions of structural connectivity, to better understand these changes in conscious awareness. In particular, we (a) examined if aspects of ongoing thought, as measured by multi-dimensional experience sampling during a sustained attention task, are associated with the white matter fibre organization of the cortex as reflected by their relative degree of anisotropic diffusion, and (b) whether these neuro-cognitive descriptions of ongoing experience are related to a more constrained measure of visual consciousness provided by the analysis of bistable perception during binocular rivalry. Individuals with greater fractional anisotropy (FA) in right hemisphere white matter regions involving the inferior fronto-occipital fasciculus, the superior longitudinal fasciculus and the cortico-spinal tract, described their ongoing thoughts as lacking external details. Subsequent analysis indicated that the combination of low FA in these right hemisphere regions, with reports of high level external details, was associated with the shortest periods of dominance during binocular rivalry. Since variation in binocular rivalry reflects differences between bottom-up and top-down influences on vision, our study suggests that reports of ongoing thoughts with vivid external details may occur when conscious precedence is given to bottom-up representation of perceptual information and that this may partly be rooted in the white matter fibre organization of the cortex.

NeuroImage: a journal of brain function 216, 116745 (2020). doi:10.1016/j.neuroimage.2020.116745, Published by Academic Press, Orlando, Fla

It is well recognized that the default mode network (DMN) is involved in states of imagination, although the cognitive processes that this association reflects are not well understood. The DMN includes many regions that function as cortical "hubs", including the posterior cingulate/retrosplenial cortex, anterior temporal lobe and the hippocampus. This suggests that the role of the DMN in cognition may reflect a process of cortical integration. In the current study we tested whether functional connectivity from uni-modal regions of cortex into the DMN is linked to features of imaginative thought. We found that strong intrinsic communication between visual and retrosplenial cortex was correlated with the degree of social thoughts about the future. Using an independent dataset, we show that the same region of retrosplenial cortex is functionally coupled to regions of primary visual cortex as well as core regions that make up the DMN. Finally, we compared the functional connectivity of the retrosplenial cortex, with a region of medial prefrontal cortex implicated in the integration of information from regions of the temporal lobe associated with future thought in a prior study. This analysis shows that the retrosplenial cortex is preferentially coupled to medial occipital, temporal lobe regions and the angular gyrus, areas linked to episodic memory, scene construction and navigation. In contrast, the medial prefrontal cortex shows preferential connectivity with motor cortex and lateral temporal and prefrontal regions implicated in language, motor processes and working memory. Together these findings suggest that integrating neural information from visual cortex into retrosplenial cortex may be important for imagining the future and may do so by creating a mental scene in which prospective simulations play out. We speculate that the role of the DMN in imagination may emerge from its capacity to bind together distributed representations from across the cortex in a coherent manner.

Neural activity within the default mode network (DMN) is widely assumed to relate to processing during off-task states, however it remains unclear whether this association emerges from a shared role in self or social content that is common in these conditions. In the current study, we examine the possibility that the role of the DMN in ongoing thought emerges from contributions to specific features of off-task experience such as self-relevant or social content. A group of participants described their experiences while performing a laboratory task over a period of days. In a different session, neural activity was measured while participants performed Self/Other judgements (e.g., Does the word ‘Honest’ apply to you (Self condition) or Barack Obama (Other condition)). Despite the prominence of social and personal content in off-task reports, there was no association with neural activity during off-task trait adjective judgements. Instead, during both Self and Other judgements we found recruitment of caudal posterior cingulate cortex—a core DMN hub—was above baseline for individuals whose laboratory experiences were characterised as detailed. These data provide little support for a role of the DMN in self or other content in the off-task state and instead suggest a role in how on-going thought is represented.

When environments lack compelling goals, humans often let their minds wander to thoughts with greater personal relevance; however, we currently do not understand how this context-dependent prioritisation process operates. Dorsolateral prefrontal cortex (DLPFC) maintains goal representations in a context-dependent manner. Here, we show this region is involved in prioritising off-task thought in an analogous way. In a whole brain analysis we established that neural activity in DLPFC is high both when ‘on-task’ under demanding conditions and ‘off-task’ in a non-demanding task. Furthermore, individuals who increase off-task thought when external demands decrease, show lower correlation between neural signals linked to external tasks and lateral regions of the DMN within DLPFC, as well as less cortical grey matter in regions sensitive to these external task relevant signals. We conclude humans prioritise daydreaming when environmental demands decrease by aligning cognition with their personal goals using DLPFC., The authors show that the dorsolateral prefrontal cortex (DLPFC) is involved in both on-task thought during increased environmental demands, and off-task thought during decreased demand–suggesting a role for the DLPFC in prioritising goals in a context-dependent manner.

Neural activity within the default mode network (DMN) is widely assumed to relate to processing during off-task states, however it remains unclear whether this association emerges from a shared role in self or social cognition. In the current study, we examine the possibility that the role of the DMN in ongoing thought emerges from contributions to specific features of off-task experience such as self-relevant or social content. A group of participants described their experiences while performing a laboratory task over a period of days. In a different session, neural activity was measured while participants performed self/other judgements. Despite the prominence of social and personal content in off-task reports, there was no association with neural activity during off-task trait adjective judgements. Instead, during both self and other judgements we found recruitment of caudal posterior cingulate cortex - a core DMN hub - was above baseline for individuals whose laboratory experiences were characterised as detailed. These data provide little support for a role of the DMN in self or other content in the off-task state and instead suggest a role in how on-going thought is represented.

Available online 30 April 2021. Brainhack is an innovative meeting format that promotes scientific collaboration and education in an open, inclusive environment. This NeuroView describes the myriad benefits for participants and the research community and how Brainhacks complement conventional formats to augment scientific progress. The present manuscript is part of a growing community effort to collate Brainhack-related insights and expertise into a Jupyter Book (http://brainhack.org/brainhack_jupyter_book/) that will serve as a centralized set of resources for the community; we acknowledge all the individuals who contributed and will make ongoing contributions to these resources. A pre-print version of the present manuscript is available as part of the Jupyter Book. Moreover, we would like to acknowledge all Brainhack organizers, supporters, presenters, and participants for their contribution to growing and maintaining this community. The benefits described in this manuscript would not be possible without them. We also thank all institutions, labs, and organizations who have helped this community grow, meet in stimulating environments, and add an excellent educational resource pool and agenda. With an expanding community, Brainhack’s support network keeps growing, and we thank all labs and individual researchers for their dedication and expertise offered to this community (see http://brainhack.org/brainhack_jupyter_book/acknowledgments.html for a full list of individual acknowledgments; an updated list will be maintained in the Jupyter Book). Grants and funding bodies: This project has received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement no. 867458 awarded to Julia M. Huntenburg; from ANR-19-DATA-0025 NeuroWebLab for Katja Heuer, Roberto Toro, and Nicholas Traut; and from NIH K00MH122372 for Stephanie Noble. The Brainhack Community member list and contributions of the different authors are detailed at http://brainhack.org/brainhack_jupyter_book/contributors.html. Our crediting system is described here: http://brainhack.org/brainhack_jupyter_book/neuroview_authorship-agreement.html.

Human cognition is flexible - drawing on both sensory input, and representations from memory, to successfully navigate complex environments. Contemporary accounts suggest this flexibility is possible because neural function is organized into a hierarchy. Neural regions are organized along a macroscale gradient, anchored at one end by unimodal systems involved with perception and action, and at the other by transmodal systems, including the default mode network, supporting cognition less directly tied to immediate stimulus input. The current study tested whether this cortical hierarchy captures modes of behaviour that depend on immediate input, as well as those that depend on representations from memory. Participants made decisions regarding the location or identity of shapes using information in the environment (0-back) or from a prior trial (1-back). Using task based imaging we established that, regardless of the nature of the decision, medial and lateral visual cortex were recruited when decisions rely on immediate input, while transmodal regions were recruited when judgments depend on information from the prior trial. Using principal components analysis, we demonstrated that shifting decision-making from perception to memory altered the focus of neural activity from unimodal to transmodal regions (and vice versa). Notably, the more pronounced these shifts in neural activity from unimodal to transmodal regions when decisions relied on memory, the more efficiently individuals performed this task. These data illustrate how the macroscale organization of neural function into a hierarchy allows cognition to rely on input, or information from memory, in a flexible and efficient manner.

Regions of transmodal cortex, in particular the default mode network (DMN), have historically been argued to serve functions unrelated to task performance, in part because of associations with naturally occurring periods of off-task thought. In contrast, contemporary views of the DMN suggest it plays an integrative role in cognition that emerges from its location at the top of a cortical hierarchy and its relative isolation from systems directly involved in perception and action. The combination of these topographical features may allow the DMN to support abstract representations derived from lower levels in the hierarchy and so reflect the broader cognitive landscape. To investigate these contrasting views of DMN function, we sampled experience as participants performed tasks varying in their working-memory load while inside an fMRI scanner. We used self-report data to establish dimensions of thought that describe levels of detail, the relationship to a task, the modality of thought, and its emotional qualities. We used representational similarity analysis to examine correspondences between patterns of neural activity and each dimension of thought. Our results were inconsistent with a task-negative view of DMN function. Distinctions between on- and off-task thought were associated with patterns of consistent neural activity in regions adjacent to unimodal cortex, including motor and premotor cortex. Detail in ongoing thought was associated with patterns of activity within the DMN during periods of working-memory maintenance. These results demonstrate a contribution of the DMN to ongoing cognition extending beyond task-unrelated processing that can include detailed experiences occurring under active task conditions.

International audience; Contemporary cognitive neuroscience recognises unconstrained processing varies across individuals, describing variation in meaningful attributes, such as intelligence. It may also have links to patterns of on-going experience. This study examined whether dimensions of population variation in different modes of unconstrained processing can be described by the associations between patterns of neural activity and self-reports of experience during the same period. We selected 258 individuals from a publicly available data set who had measures of resting-state functional magnetic resonance imaging, and self-reports of experience during the scan. We used machine learning to determine patterns of association between the neural and self-reported data, finding variation along four dimensions. ‘Purposeful’ experiences were associated with lower connectivity -in particular default mode and limbic networks were less correlated with attention and sensorimotor networks. ‘Emotional’ experiences were associated with higher connectivity, especially between limbic and ventral attention networks. Experiences focused on themes of ‘personal importance’ were associated with reduced functional connectivity within attention and control systems. Finally, visual experiences were associated with stronger connectivity between visual and other networks, in particular the limbic system. Some of these patterns had contrasting links with cognitive function as assessed in a separate laboratory session -purposeful thinking was linked to greater intelligence and better abstract reasoning, while a focus on personal importance had the opposite relationship. Together these findings are consistent with an emerging literature on unconstrained states and also underlines that these states are heterogeneous, with distinct modes of population variation reflecting the interplay of different large-scale networks.

The tendency for the mind to wander to concerns other than the task at hand is a fundamental feature of human cognition, yet the consequences of variations in its experiential content for psychological functioning are not well understood. Here, we adopted multivariate pattern analysis to simultaneously decompose experience-sampling data and neural functional-connectivity data, which revealed dimensions that simultaneously describe individual variation in self-reported experience and default-mode-network connectivity. We identified dimensions corresponding to traits of positive-habitual thoughts and spontaneous task-unrelated thoughts. These dimensions were uniquely related to aspects of cognition, such as executive control and the ability to generate information in a creative fashion, and independently distinguished well-being measures. These data provide the most convincing evidence to date for an ontological view of the mind-wandering state as encompassing a broad range of different experiences and show that this heterogeneity underlies mind wandering’s complex relationship to psychological functioning.

We often think about people, places and events that are outside of our immediate environment. Although prior studies have explored how we can reduce the occurrence of these experiences, the neurocognitive process through which they are produced are less understood. The current study builds on developmental and evolutionary evidence that language helps organise and express our thoughts. Behaviorally, we found the occurrence of task unrelated thought (TUT) in easy situations was associated with thinking in words. Using experience sampling data, in combination with online measures of neural function, we established that activity in a region of anterior cingulate cortex / medial-prefrontal cortex (mPFC) tracked with changes in the expression of TUT. This region is at the intersection of two mPFC clusters identified through their association with variation in aspects of spontaneous thought: thinking in words (dorsal) and mental time travel (ventral). Finally, using meta-analytic decoding we confirmed the dorsal/ventral distinction within mPFC corresponding to a functional difference between domains linked to language and meaning and those linked to memory and scene construction. This evidence suggests a role for mPFC in the expression of TUT that may emerge from interactions with distributed neural signals reflecting processes such as language and memory.

The default mode network supports a variety of mental operations such as semantic processing, episodic memory retrieval, mental time travel and mind-wandering, yet the commonalities between these functions remains unclear. One possibility is that the default mode network supports cognition that is independent of the immediate environment; alternatively or additionally, it might support higher-order conceptual representations that draw together multiple features. We tested these accounts using a novel paradigm that contrasted decisions based on perceptual and conceptual features, while separately manipulating whether these decisions were driven by features in the environment or recovered from memory. Task-based fMRI identified regions that responded when stimulus independence and semantic retrieval were combined: these included left and right angular gyri and left middle temporal gyrus. Although these sites were within the default mode network, they showed a stronger response to demanding memory judgements than to an easier perceptual task, contrary to the view that they support automatic aspects of cognition. In a subsequent analysis, we showed that these regions were located at the extreme end of a macroscale gradient, which describes gradual transitions from sensorimotor to transmodal cortex. This shift in the focus of neural activity towards transmodal default mode regions might reflect isolation from specific sensory inputs, both when decisions are guided by conceptual as opposed to perceptual features and when cognitive states are generated in the absence of input.

Trying to focus on a piece of text and keep unrelated thoughts at bay can be a surprisingly futile experience. The current study explored the effects of different instructions on participants’ capacity to control their mind-wandering and maximize reading comprehension, while reading. Participants were instructed to (a) enhance focus on what was read (external) or (b) enhance meta-awareness of mind-wandering (internal). To understand when these strategies were important, we induced a state of self-focus in half of our participants at the beginning of the experiment. Results replicated the negative association between mind-wandering and comprehension and demonstrated that both internal and external instructions impacted on the efficiency of reading following a period of induced self-focus. Techniques that foster meta-awareness improved task focus but did so at the detriment of reading comprehension, while promoting a deeper engagement while reading improved comprehension with no changes in reported mind-wandering. These data provide insight into how we can control mind-wandering and improve comprehension, and they underline that a state of self-focus is a condition under which they should be employed. Furthermore, these data support component process models that propose that the self-referent mental contents that arise during mind-wandering are distinguishable from those processes that interfere with comprehension.