Nijhof AD, Willems RM (2015) Simulating Fiction: Individual Differences in Literature Comprehension Revealed with fMRI. PLoS ONE 10(2): e0116492. doi:10.1371/journal.pone.0116492
Abstract: When we read literary fiction, we are transported to fictional places, and we feel and think along with the characters. Despite the importance of narrative in adult life and during development, the neurocognitive mechanisms underlying fiction comprehension are unclear. We used functional magnetic resonance imaging (fMRI) to investigate how individuals differently employ neural networks important for understanding others’ beliefs and intentions (mentalizing), and for sensori-motor simulation while listening to excerpts from literary novels. Localizer tasks were used to localize both the cortical motor network and the mentalizing network in participants after they listened to excerpts from literary novels. Results show that participants who had high activation in anterior medial prefrontal cortex (aMPFC; part of the mentalizing network) when listening to mentalizing content of literary fiction, had lower motor cortex activity when they listened to action-related content of the story, and vice versa. This qualifies how people differ in their engagement with fiction: some people are mostly drawn into a story by mentalizing about the thoughts and beliefs of others, whereas others engage in literature by simulating more concrete events such as actions. This study provides on-line neural evidence for the existence of qualitatively different styles of moving into literary worlds, and adds to a growing body of literature showing the potential to study narrative comprehension with neuroimaging methods.
Narratives play an important role in human life, and it is more and more acknowledged that fiction is a powerful player in human development as well as in adulthood (e.g. [1,2,3]). Despite its importance, it is largely unknown what the brain networks are that support our unique ability to move into a fiction world. While it is uncontroversial that people are moved into fiction worlds [4,5], it is unclear how readers do this. People differ greatly in how they engage in fiction (e.g. [6,7–11]), but the neurocognitive mechanisms behind narrative engagement remain unclear (see  for related work on theatre). Here we use neuroimaging to investigate individual differences during the comprehension of literary fiction stories.
One way in which participants engage with stories, is via simulation of the story’s content. Recent philosophical and neuroscientific evidence shows that it is important to distinguish at least two neurocognitively distinct components of simulation when considering the understanding of narratives (e.g. ). First, sensori-motor simulation is evidenced by activation of motor and visual cortices when people comprehend language related to actions and scenery [14–17]. The second component relates to our ability to understand thoughts, intentions and beliefs of others, sometimes called mentalizing . The distinction between these two components important for fiction understanding is theoretically motivated (e.g. ), and supported by neural findings (e.g. [19,20]).
In this study participants listened to excerpts (4 to 8 minutes long) from literary novels, while neural activity was measured across the whole brain by means of fMRI. We chose to use listening rather than word-by-word reading, because relatively long fragments are used and therefore listening is expected to be the most convenient option for participants in the scanner. Supposedly, this would not result in crucial differences in terms of the mental simulation they employ. Previous studies did find differences in brain activity between listening and reading (with more individual differences in activity for reading), but also several core regions shared between modalities . More importantly for the purposes of the present study, it has been shown that regions involved in mentalizing  and action understanding  are activated independent of presentation modality. It is an open question whether mentalizing differs during reading or listening to narratives, but based on the previous literature we expect the two modalities to engage overlapping neural correlates.
Stories were tagged for motor (‘action’) and mentalizing content, and memory for the stories was debriefed afterwards. Brain regions known to be involved in the two kinds of simulation were localized with standardized localizer tasks. Target regions were left and right motor regions for sensori-motor simulation , and anterior medial prefrontal cortex (amPFC), right temporoparietal junction (rTPJ) and precuneus for mentalizing [19,25] (see methods section). Importantly, measurement of brain activity during story comprehension was done on-line, and without additional tasks for the listener.
Some recent neuroimaging studies relate to the issue of mentalizing and sensori-motor simulation during the comprehension of narratives. For instance, Wallentin and colleagues showed that part of the visual cortex which is sensitive to perceiving visual motion, is also activated when participants heard pieces describing movement in a retelling of ‘The Ugly Duckling’ . Similarly, Speer and colleagues showed that parts of short children’s stories containing action descriptions activated the motor cortex . In an interesting recent approach, Altmann and colleagues presented short stories (around 40 words per story) to participants. Stories were either labeled as fact (describing an event that actually took place) or as fiction. Most interesting for the current approach was that stories that were labeled as fiction led to stronger activation in medial prefrontal cortex (among other regions), whereas labeling stories as describing actual facts led to higher activation levels in the premotor cortex (again, amongst other regions) (; see also [14,28]).
In this study we follow up on this previous work by using more extended (i.e. longer) excerpts from literary fiction, written for adults, in order to give participants an experience of engaging with fiction that is relatively close to their real-world experience. We have a special focus on individual differences in simulation and mentalizing during narrative comprehension. Previous work has found that participants differ in how much they engage parts of the mentalizing system during the reading of texts labeled as fiction , as well as that participants differ in how much they engage in sensori-motor simulation during language comprehension [29,30]. Here we combine these two to see how participants differ in their engagement of these two important subprocesses of narrative comprehension while listening to natural, unmodified literary fiction.
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