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Neuroimaging Studies of the Default Mode Netword

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In the past couple of blog posts, I have detailed various behavioral studies that have conclusively supported our shared conscious experience of mind wandering. When our minds are redirected from the external environment to internal mentations, our accuracy at sustained attention tasks decreases and we can read entire pages of a book without actually comprehending of it. But this same process allows creative incubation and goal-directed planning for the future. Like planning for the future, most of our conscious experience of mind wandering is spent ruminating about our recent past and imaging our possible futures. At the neural level, what allows us to create these spontaneously fullblown, movie-length, extremely detailed daydreams without any external stimuli guiding our thoughts? Thankfully, with the advancements in neuroimaging technology, we are able to study the neural underpinnings of not only task-related behavior, but also the conscious experience of mind wandering.

From previous brain imaging research, we know that certain brain regions are active during a task (task-positive brain responses) while others show less activation during a task (task-negative brain responses). Not only are certain brain regions active during a task, but their activations are also correlated with one another. In 2005, Fox et al. wanted to know whether or not these different brain regions, both task-positive and task-negative, are correlated with one another in the resting brain. If so, this would support the idea that task-related functionality is represented intrinsically in the brain, meaning that the human brain is organized into two anticorrelated functional networks.

In order to test this theory, Fox et al. analyzed resting state fMRI data for ten normal individuals and analyzed correlations within and between three regions of interest known to be task-positive (intraparietal sulcus (IPS), frontal eye field, and middle temporal region) and three regions of interest known to be task-negative (medial prefrontal cortex (mPFC), posterior cingulate/precuneus (PCC), and lateral parietal cortex). Imaging data at the individual level when the task-positive PCC was used as a seed region showed a correlation with the task-positive region mPFC and an anticorrelation with the task-negative region IPS (shown below). The time course data presented below the brain images demonstrates that both relationships are stable over time as well.

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The z-scored group data provides even more compelling evidence for the correlation of brain regions within both the task-positive and task-negative brain networks, and shows that these networks are anticorrelated when comparing between the regions. The imaging data below shows how the task-positive seed regions look almost identical (left panel) as well as the task-negative seed regions (right panel). However, the most compelling aspect of this analysis is the conjunction map, which shows how regions within each network are correlated, while the networks are anticorrelated with one another.

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So it is clear that there are two distinct brain networks associated with increases and decreases in endogenous attentional demand whose differences are highlighted in resting state image data. The authors argue that these networks are intrinsic, as they were able to reproduce their results for three different resting states (fixation, eyes closed, and eyes opened). So what do we make of this anticorrelation? According to the time course data, it appears as though during rest, the two networks simultaneously oscillate between activation and deactivation. It is difficult to tell if this relationship is merely correlational or if the networks are influencing each other. It would have been interesting if the authors could have performed a time lag analysis in order to determine if when one network activates, it has an almost macro level lateral inhibition effect on the other network, resulting in its deactivation. This might shed light on the functionality of these two brain networks anticorrelated relationship. One could utilize TMS during the same resting state study in order to stimulate one brain region within the task-positive network to see if this has deactivating effects on the task-negative regions. Fox et al. argue that there study provides evidence that rejects the notion of a resource competition theory of mind wandering and sustained attention because there are two distinct networks, however, their study utilized a resting state paradigm in which attentional resources are not needed at all and we have no idea what the conscious experience of each subject was during the scan. In order to better understand the relationship between these two supposedly anticorrelated networks, we need to look at studies that require the subject to give behavioral responses as well as report their conscious experiences. In 2007, Mason et al. utilized fMRI to demonstrate default mode network (task-negative regions, DMN) activation during periods of mind wandering. By training subjects on their task so that they were proficient enough to mindlessly complete it (ie mind wander), Mason et al. were able to demonstrate a positive correlation between stimulus independent thought frequency and DMN activation. Additionally, DMN activation accounted for individual differences in propensity to mind wander (as measured by the Imaginal Process Inventory). Mason et al.s study supports the notion that the task-negative regions of the DMN serve as the neural support for mind wandering. While this is compelling, it is still possible that the DMN could also support external attention in addition to internal mentation. Studies with contrasting findings to Mason et al. have shown that certain regions of the DMN are activated during periods of broad awareness of our external world, possibly allowing us to be ready to respond to unpredictable stimuli. This theory does not contradict Fox et al.s findings, as participants were in a resting state where their awareness could easily broaden due to lack of any task to focus on. In order to investigate the validity of both claims, Andrews-Hanna et al. conducted an imaging study in 2010 in which they controlled for external stimuli across conditions while manipulating only expectations of task in order to vary spontaneous cognition (as operationalized by direction and scope of attention). They utilized a vigilance task in which subjects detected either a peripheral stimulus (broad monitoring), a central stimulus (foveal monitoring), or simply fixated on a central point (resting state passive, essentially). The clever part of this paradigm is that they only analyzed data from trials with equal stimuli across conditions. In order to measure conscious experience, an independent group was probed for task relevant (fixation point and additional stimuli) or task irrelevant thoughts (past and future) while in a mock scanner. The authors
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reported data, which they argued, supported the notion that the DMN is involved with internal mentation rather than broad external monitoring due to the DMN being most active during the passive conditions. They also demonstrated equivalent activations for broad and focal conditions, both being lower than passive, showing that the DMN is not responsible specifically for broad external monitoring (shown below).

However, their paradigm has some key flaws that need to be addressed. Firstly, there were no online measures of conscious experience during the scanning, instead Andrews-Hanna et al. extrapolated SIT measurements from an independent group of subjects. Even more problematic is that stimulus independent thought can be both task-relevant and task-irrelevant. Meaning, one can think about the task at hand and not have this thought generated by or regarding the stimulus being presented. In order to address this problem, Stawarczyk et al. (2011) conducted a study in which they probed participants online while in the scanner taking into account both task relatedness and stimulus dependence as classifications for conscious experience (see divisions of four categories below).

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By taking into account both of these possible conscious experience dimensions, Stawarczyk et al. were able to demonstrate that specific regions within the DMN are responsible for mind wandering (or internal mentation) while others are responsible for unfocused external attention. For example, a main effect was shown for task relatedness when looking at differences in activation between a rostral portion of the mPFC versus a more caudal but adjacent portion of it (seen below). Even more striking is that an additive effect was shown for taskrelatedness and stimulus-dependency for the mPFC and the PCC/precuneus, meaning that DMN activity was highest for mind wandering conditions, followed by external distractions and task-related interferences, and least active for taskfocused conditions.

This study reveals the complex nature of the DMN, which has potential to help clear up previous disputes about the DMN and mind wandering. For instance, even though mind wandering episodes engage both executive networks and episodic networks, this may be explained by internalized autobiographical planning. Allowing for a mixture of categories of conscious thought, we are better able to reconcile these findings. One thing is for sure, studying conscious experience cannot be oversimplified to on task versus off task attention. Our conscious experience is wrought with complexities, and utilizing paradigms that allow for better categorization of our thoughts, like Stawarczyk et al. 2011, will better illuminate the role of the DMN in external and internal attention.

References Fox, M. D., A. Z. Snyder, et al. (2005). The human brain is intrinsically organized into dynamic, anticorrelated functional networks. Proc Natl Acad Sci U S A 102(27): 96739678. Mason, M. F., M. I. Norton, et al. (2007). Wandering minds: the default network and stimulus-independent thought. Science 315(5810): 393-395. Andrews-Hanna, J. R., J. S. Reidler, et al. (2010). Evidence for the default networks role in spontaneous cognition. J Neurophysiol 104(1): 322-335. Stawarczyk, D., S. Majerus, et al. (2011). Neural correlates of ongoing conscious experience: both task-unrelatedness and stimulus-independence are related to default
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network activity. PLoS One 6(2): e16997

Posted on March 26, 2013. | Leave a comment

The Benefits and Disadvantages of Mind Wandering

Sort of in the same vein as dreaming, because mind wandering is a universally experienced mental state, we would think that it must have some evolutionary benefit to the human species. However, most of the research conducted on mind wandering highlights the consequences associated with it. In fact, all of my blog posts have focused on mind wandering as getting off task and not paying attention, thus leading to performance decrements. Sure mind wandering can definitely hurt our performance when doing homework, reading for class, or listening to a crucial lecture. But we also mind wander when we are not completing a task, therefore there must be some functionality associated with this ever present mental state. Before speculating about the positive effects of mind wandering, we must remember how detrimental it can be when performing a task. McVay and Kane, the same authors who assert the executive failure theory of mind wandering, demonstrated in their 2012 paper Why Does Working Memory Capacity Predict Variation in Reading Comprehension? On the Influence of Mind Wandering and Executive Attention, that the ability to control our attention, thus disabling our minds from wandering, mediates the strong correlation between working memory capacity (WMC) and reading comprehension. In this paper, McVay and Kane hypothesized that if individual differences in task unrelated thoughts (TUTs) are consistent across attention and reading tasks and if the propensity to mind wander predicts comprehension, then this would provide stronger evidence for inefficient attentional control as a cause of poor reading. Secondly, the executive component of WMC, not the memory component, will predict reading comprehension. And lastly, TUT rate should mediate the relationship between WMC and reading comprehension. Participants participated in three sessions during which they completed working memory tasks, attentional-control tasks, and reading comprehension tasks. After several analyses utilizing latent-variable structural-equation-model approaches, McVay and Kane found support for their three hypotheses in their data. Ultimately they reconciled their views and findings with Smallwoods revised proposal (discussed in the previous blogpost), positing that during cognitive activities, failure to restrict attention to goal-related representations will result in mind wandering intrusions leading to comprehension errors. Sometimes mind wandering can not only cause us to forget the information we were supposed to be attending to during the daydream episode, but also the information we had previously been encoding. In their article Remembering to Forget: The Amnesic Effect of Daydreaming, Delaney et al. (2010) posit that mind wandering can disrupt the encoding of information previously being attended to. For example, when listening to a boring lecture we might attend to the first twenty minutes of the talk, however our mind may start to wander during that last forty minutes. Later on, we certainly wont know what the lecturer was
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talking about during the last forty minutes, but we might also forget the first twenty. How can this be? Previous research has shown that thoughts similar in content to daydreams can impair memory for newly learned information. In this study, they utilized a diversion paradigm in which subjects learned a list of words, were instructed to divert their thinking, and then study a second list. During the subsequent free recall test, participants in the experimental condition (diversionary thoughts) forgot more of the first list than subjects in the control group. The authors posit that items from list one are harder to recall in the new mental landscape contextualized by the diversionary thought. This relates to Tulvings Encoding Specificity Principle in that the context of recall does not match the context at encoding. Thus, they hypothesized that the extent to which one forgets list one depends on how different the current context is relative to the imagined one. In order to test this prediction against the amnesic effect of mind wandering, Delaney et al. conducted two different diversion paradigms. In the first, they were able to demonstrate that subjects who were asked to think about their parents home versus their own home forgot more of list one, presumably because the home of a college students parents is thought to be more temporally distant than their current home. In the second, they showed that subjects who thought about a past international trip forgot more of list one than those who thought of a domestic vacation, supposedly because the international destination is more spatially distant than a domestic one (results below).

Delaney et al. admit that their conditions are not pure, rather they are a mixture of both temporal and spatial mental proximities. This reflected in the incongruous finding between conditions, that near-change subjects differed from controls during the spatial manipulation but not during the temporal one. However, they were able to establish negative correlations for both miles and weeks with proportion of list one recalled (results below).

While this study supports the notion that task unrelated thoughts may disrupt the
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subsequent recall of a previously encoded word list, it does not accurately depict the effects of mind wandering. More specifically, daydreaming is not a consciously directed thought like the diversionary thoughts utilized in this experiment. One thing that Schooler and McVay agree on is that mind wandering is automatically and unconsciously initiated, therefore it seems like the diversionary thoughts Delaney et al. told their subjects to explicitly think about do not qualify as daydreams. In order to avoid this artificiality, many researchers opt to choose more natural data collection methods. For example, Killingsworth and Gilberts 2010 Science paper A Wandering Mind Is an Unhappy Mind, utilized an awesome application for the iPhone that contacts subjects at random throughout the day asking them How are you feeling right now?, What are you doing right now?, and Are you thinking about something other than what youre currently doing?. This way, Killingsworth and Gilbert were able to collect copious amounts of natural data from various types of individuals (2,250 subjects!). They found that people were less happy when their minds were wandering than when they were not. Surprisingly, this was true of all types of activities even the least enjoyable, even though people were most likely to wander to positive thoughts. In order to reject the notion that peoples minds wandered when they were upset, the researchers conducted time-lag analyses that strongly suggest the opposite causal relationship, people were less happy because they were mind wandering. Because I truly want to believe that daydreaming of fond memories from the past and imagining exciting opportunities that might come to fruition does not cause an individual to be less happy, I am skeptical about the findings presented in this article. First, I wonder if the authors conducted baseline propensities to mind wander for all of their subjects. Maybe there is a strong correlation between likelihood of daydreaming and likelihood of not being happy, but not necessarily a causal relationship. Nevertheless, in light of these findings it is definitely important that we give serious consideration to the assertion that staying present-oriented and not ruminating on the past and future results in a more happy state of being. Even though mind wandering during reading can result in comprehension deficiencies, and mind wandering throughout the day can lead to an emotional state that is less happy, there still exists some potential benefits of mind wandering. While Delaney et al. assert that the diversion paradigm is representative of daydreaming, I think it is merely demonstrating a handicap to rehearsal of a word list as a result of an experimenter-induced intrusive thought. Conversely, true daydreaming may have beneficial effects on higher order cognitive thinking. In their paper Inspired by Distraction: Mind Wandering Facilitates Creative Incubation, Baird et al. (2012) demonstrated that creativity could be enhanced for finding solutions to a higher order cognitive problem after participants engaged in mind wandering. Participants were asked questions like come up with as many unusual uses of a brick as you can in a minute, then given an undemanding task (assumed to result in the highest frequency of mind wandering), a demanding task, rest, or no break. Participants in the undemanding task condition (which was shown to produce more task unrelated thoughts after a retroactive questionnaire) improved the most on the unusual uses task (results shown below).

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Mind wandering did not, however, improve creative thinking in general, shown by a lack of condition differences when given a new problem. Therefore, mind wandering helped subjects in terms of an incubation effect, allowing them to unconsciously process associations in terms of the default network. Thus, mind wandering may help increase creative inspiration, leading to the crucial ahha moments of insight that one experiences when not explicitly thinking about the problem at hand. It appears that mind wandering has both advantages and disadvantages to cognitive processing then depending on when the daydreaming takes place in relation to the task at hand. It seems like mind wandering might also play a role in episodic memory consolidation, very much like the dreaming state of sleep, which would make sense intuitively as we call mind wandering daydreaming. As long as our attention is not being demanded of by external stimuli, mind wandering seems to be a beneficial process for creative thinking and problem solving. Of course we must strike a balance, as Killingsworth and Gilbert warn in their paper, between living in the now and living in our past and future oriented daydreams. References Killingsworth, M. A. and D. T. Gilbert (2010). A wandering mind is an unhappy mind. Science 330(6006): 932. McVay, J. C. and M. J. Kane (2011). Why does working memory capacity predict variation in reading comprehension? On the influence of mind wandering and executive attention. J Exp Psychol Gen. Delaney, P. F., L. Sahakyan, et al. (2010). Remembering to forget: The amnesic effect of daydreaming. Psychol Sci 21(7): 1036-1042. Baird, B., J. Smallwood, et al. (2012). Inspired by distraction: Mind Wandering facilitates creative incubation. Psychol Sci.
Posted on March 5, 2013. | Leave a comment

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