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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.

The hippocampus contributes to episodic, spatial and semantic aspects of memory, yet individual differences within and between these functions are not well-understood. In 136 healthy individuals, we investigated whether these differences reflect variation in the strength of connections between functionally-specialised segments of the hippocampus and diverse cortical regions that participate in different aspects of memory. Better topographical memory was associated with stronger connectivity between lingual gyrus and left anterior, rather than posterior, hippocampus. Better semantic memory was associated with increased connectivity between the intracalcarine/cuneus and left, rather than right, posterior hippocampus. Notably, we observed a double dissociation between semantic and topographical memory: better semantic memory was associated with stronger connectivity between left temporoparietal cortex and left anterior hippocampus, while better topographic memory was linked to stronger connectivity with right anterior hippocampus. Together these data support a division-of-labour account of hippocampal functioning: at the population level, differences in connectivity across the hippocampus reflect functional specialisation for different facets of memory, while variation in these connectivity patterns across individuals is associated with differences in the capacity to retrieve different types of information. In particular, within-hemisphere connectivity between hippocampus and left temporoparietal cortex supports conceptual processing at the expense of spatial ability.

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.

The ability to visualise different people and places in imagination is a core element of human cognition. Contemporary neuroimaging research highlights regions of primary visual cortex as important in instantiating images in the mind’s eye. Here we combine task-based functional magnetic resonance imaging with measures of intrinsic brain organisation to show the ventral attention, or salience, network, also plays an important role in visual imagination. In a task-based study we replicated prior observations of regions of medial occipital cortex, including Brodmann Area 17, showing greater neural activity during imagination than when perceiving similar stimuli. In addition, we found regions of the ventral attention network, and in particular right dorsolateral prefrontal cortex (BA 9/46), were activated by the same acts of imagination. In a subsequent resting-state study, we demonstrated medial occipital cortex regions form a functional network with regions of the ventral attention network that were also active during imagination. Furthermore, this pattern was maximised in an overlapping region of right dorsolateral prefrontal cortex for individuals whose self-reported imagery had stronger negative affect. Together these data suggest visual imagination relies on functional interactions between regions of primary visual cortex with the ventral attention system, and in particular, with a region of right dorsolateral prefrontal cortex.

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.

Theoretical accounts of face processing often emphasise feature shapes as the primary visual cue to the recognition of facial expressions. However, changes in facial expression also affect the surface properties of the face. In this study, we investigated whether this surface information can also be used in the recognition of facial expression. First, participants identified facial expressions (fear, anger, disgust, sadness, happiness) from images that were manipulated such that they varied mainly in shape or mainly in surface properties. We found that the categorization of facial expression is possible in either type of image, but that different expressions are relatively dependent on surface or shape properties. Next, we investigated the relative contributions of shape and surface information to the categorization of facial expressions. This employed a complementary method that involved combining the surface properties of one expression with the shape properties from a different expression. Our results showed that the categorization of facial expressions in these hybrid images was equally dependent on the surface and shape properties of the image. Together, these findings provide a direct demonstration that both feature shape and surface information make significant contributions to the recognition of facial expressions.

Human cognition is not always tethered to events in the external world. Laboratory and real world experience sampling studies reveal that attention is often devoted to self-generated mental content rather than to events taking place in the immediate environment. Recent studies have begun to explicitly examine the consistency between states of off-task thought in the laboratory and in daily life, highlighting differences in the psychological correlates of these states across the two contexts. Our study used neuroimaging to further understand the generalizability of off-task thought across laboratory and daily life contexts. We examined (1) whether context (daily life versus laboratory) impacts on individuals’ off-task thought patterns and whether individual variations in these patterns are correlated across contexts; (2) whether neural correlates for the patterns of off-task thoughts in the laboratory show similarities with those thoughts in daily life, in particular, whether differences in cortical grey matter associated with detail and off-task thoughts in the para-hippocampus, identified in a prior study on laboratory thoughts, were apparent in real life thought patterns. We also measured neural responses to common real-world stimuli (faces and scenes) and examined how neural responses to these stimuli were related to experiences in the laboratory and in daily life - finding evidence of both similarities and differences. There were consistent patterns of off-task thoughts reported across the two contexts, and both patterns had a commensurate relationship with medial temporal lobe architecture. However, compared to real world off-task thoughts, those in the laboratory focused more on social content and showed a stronger correlation with neural activity when viewing faces compared to scenes. Overall our results show that off-task thought patterns have broad similarities in the laboratory and in daily life, and the apparent differences may be, in part, driven by the richer environmental context in the real world. More generally, our findings are broadly consistent with emerging evidence that shows off-task thoughts emerge through the prioritisation of information that has greater personal relevance than events in the here and now., Highlights • Laboratory and daily life patterns of thought show both similarities and differences. • Thoughts from both contexts have similar associations with medial temporal lobe grey matter. • Laboratory off-task thoughts have stronger emphasis on content related to other people. • Laboratory off-task thoughts are linked to greater neural activity when viewing faces.

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.

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.

Reversing the luminance values of a face (contrast negation) is known to disrupt recognition. However, the effects of contrast negation are attenuated in chimeric images, in which the eye region is returned to positive contrast (S. Gilad, M. Meng, & P. Sinha, 2009, Role of ordinal contrast relationships in face encoding, Proceedings of the National Academy of Sciences, USA, Vol. 106, pp. 5353–5358). Here, we probe further the importance of the eye region for the representation of facial identity. In the first experiment, we asked to what extent the chimeric benefit is specific to the eye region. Our results showed a benefit for including a positive eye region in a contrast negated face, whereas chimeric faces in which only the forehead, nose, or mouth regions were returned to positive contrast did not significantly improve recognition. In Experiment 2, we confirmed that the presence of positive contrast eyes alone does not account for the improved recognition of chimeric face images. Rather, it is the integration of information from the positive contrast eye region and the surrounding negative contrast face that is essential for the chimeric benefit. In Experiment 3, we demonstrated that the chimeric benefit is dependent on a holistic representation of the face. Finally, in Experiment 4, we showed that the positive contrast eye region needs to match the identity of the contrast negated part of the image for the chimera benefit to occur. Together, these results show the importance of the eye region for holistic representations of facial identity.

The ability to perceive facial expressions of emotion is essential for effective social communication. We investigated how the perception of facial expression emerges from the image properties that convey this important social signal, and how neural responses in face-selective brain regions might track these properties. To do this, we measured the perceptual similarity between expressions of basic emotions, and investigated how this is reflected in image measures and in the neural response of different face-selective regions. We show that the perceptual similarity of different facial expressions (fear, anger, disgust, sadness, happiness) can be predicted by both surface and feature shape information in the image. Using block design fMRI, we found that the perceptual similarity of expressions could also be predicted from the patterns of neural response in the face-selective posterior superior temporal sulcus (STS), but not in the fusiform face area (FFA). These results show that the perception of facial expression is dependent on the shape and surface properties of the image and on the activity of specific face-selective regions.

The amygdala is known to play an important role in the response to facial expressions that convey fear. However, it remains unclear whether the amygdala’s response to fear reflects its role in the interpretation of danger and threat, or whether it is to some extent activated by all facial expressions of emotion. Previous attempts to address this issue using neuroimaging have been confounded by differences in the use of control stimuli across studies. Here, we address this issue using a block design functional magnetic resonance imaging paradigm, in which we compared the response to face images posing expressions of fear, anger, happiness, disgust and sadness with a range of control conditions. The responses in the amygdala to different facial expressions were compared with the responses to a non-face condition (buildings), to mildly happy faces and to neutral faces. Results showed that only fear and anger elicited significantly greater responses compared with the control conditions involving faces. Overall, these findings are consistent with the role of the amygdala in processing threat, rather than in the processing of all facial expressions of emotion, and demonstrate the critical importance of the choice of comparison condition to the pattern of results.