Your Brain on Story

Stories saturate all aspects of human life, spinning tales of love, power, struggles, and death. For over 100,000 years, human cultures have relied on myths, fables, legends, and folktales to make meaning of the world around them (Haven, 2007, p. 3). Stories evolved as an automatic and subconscious way to build social bonds and share lifesaving information by simulating intense experiences without the firsthand danger (Cron, 2012, p. 9). This evolutionary advantage is reinforced today by story’s endless use throughout childhood. Children still develop deep, long-lasting emotional attachments to stories told across different media; they express strong feelings, ask for repeat tellings, and reenact them in pretend play (Alexandar et al., 2001). Because neural networks can be rewired based on repeated patterns of activation, story structure is continuously strengthened and reinforced in the brain by story’s omnipresence in daily life.

While people have tried to explain the power of stories for centuries, only recently have neuroimaging and other physiological tools begun to unravel how stories activate and influence the brain. In particular, functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) provide helpful proxies for measuring metabolic brain activity. fMRI measures changes in the ratio of oxygenated to deoxygenated blood while PET measures the flow of blood to brain areas. Based on existing neuroimaging and physiological research, a neuroscientific theoretical framework of story can be constructed to understand and predict how stories affect people.

This story framework is influenced narrative transportation theory (Gerrig, 1993; Green & Brock, 2000) which argues that stories mentally transport people into the storyworld. Storyreceivers “enter, Alice-like, through a looking glass into an imagined world in which the self can become engaged and potentially transformed” (Oatley, 2013, p. 51). At the end, the storyreceiver returns from the storyworld changed in some way (Nell, 2013). With this influence, the story framework presented here draws on brain activity to explain how stories engage people, provide a round-trip ticket into the storyworld, and leave a lasting impression.

Captivating Characters

The story framework requires emotionally-engaging characters with knowable mental processes. Neuroimaging studies highlight several brain areas that contribute to mental attributions by assigning intentions and motive to characters. In addition, the motor cortex simulates characters’ actions to understand their intentions and motivations.

Theory of Mind

Inferring mental states in oneself and others, known as Theory of Mind, allows people to explain and predict others’ behavior (Mar, 2004). These mental inferences play a role in story comprehension because they allow the storyreceiver to better emotionally relate to the characters and understand how they might interact with them in real life (Mar, 2011).

Emotions. Because stories simulate the problems of human life, characters allow the storyreceiver to experience another’s emotions. In turn, the storyreceiver’s emotions are transformed, deepened, and stretched by the character’s emotions. Paul Zak quips that “this is why your palms sweat when James Bond dodges bullets. And why you stifle a sniffle when Bambi’s mother dies” (2015, p. 7). fMRI studies reveal that stories with mentalized characters activate the temporal poles, which are associated with social and emotional processing for memory (Nakumura et al., 2001; Snowden et al., 2004). In an fMRI study (Gallagher et al., 2000), participants read and viewed Theory of Mind stories and physical stories. Theory of Mind stories (“the burglar turns around, sees the policeman, and gives himself up”) required inferring mental states of characters. Similarly-plotted physical stories (“he skillfully picks the lock on the shop door”) did not require mental inferences. Researchers discovered that Theory of Mind stories bilaterally activated the temporal poles significantly more than physical stories. Another fMRI study (Berthoz et al., 2002) revealed bilateral temporal pole activity when characters violated social norms. These studies suggest that story framework requires mentalized characters to engage the brain emotionally; creating those emotional ties makes stories more memorable.

The amygdala is also implicated in stories involving emotionally-engaging, mentalized characters. An fMRI study (Ferstl et al., 2014) found that emotional characterizations, presented as descriptions of protagonists’ feelings, activated the amygdaloid complex, which is related to emotional processing, decision-making, and interpersonal interactions. In another study (AbdulSabur et al., 2014), participants told and listened to stories related to sets of three picture cards. Each story expressed a character’s emotions. For instance, one story featured a fisherman catching a lobster, looking it in the eyes while holding it over a pot of boiling water, and then throwing it back in the ocean. fMRI and PET scans revealed that the amygdala was bilaterally activated during the stories to process characters’ emotions.

Intentions and Motivations. Stories engage people to understand characters’ intentions and motivations, fulfilling an evolutionary craving to understand others and how they will behave. When Heider and Simmel (1944) showed participants a film of geometric shapes moving around randomly, most participants invented a story by assigning intent and motive to the shapes. A recent fMRI study (Nguyen et al., 2019) updated the classic experiment by having participants either watch an animated movie depicting a story through geometric shape movement or listen to a narrated version of the story. Both conditions had similar neural activity, reinforcing the brain’s wiring for attributing mental states to characters — whether human or not — and drawing inferences from whatever information is available.

As part of their evolutionary advantage, stories help decipher and predict others’ behavior from incomplete information. The left medial frontal gyrus, a ridge on the surface of the goal-centric frontal lobe, helps reduce any uncertainty about the intentions behind characters’ actions. In a PET study (Fletcher et al., 1995), Theory of Mind stories activated the left medial frontal gyrus while physical stories did not. In addition, the medial prefrontal cortex also helps clarify the intentions or motivations behind a character’s actions. Also located in the goal-centric frontal lobe, the medial prefrontal cortex plays a role in motivation, orientation, attention, and recognizing intentional actions (AbdulSabur et al., 2014). It uses self-referential information to mentalize about characters. The dorsal portion of the medial prefrontal cortex is especially engaged in assessing characters’ motivations. In an fMRI study (Jenkins & Mitchell, 2010), participants read stories with ambiguous story inferences (“her neighbor told her she might want to take a look at her tulip beds”) and stories with unambiguous story inferences (“the tulips next to her office still have not flowered yet”). fMRI revealed that the ventral and dorsal parts of the medial prefrontal cortex respond more to ambiguous than unambiguous story inferences, activated to explain characters’ intentions and motivations when not self-evident from their actions.

In the social norms fMRI study (Berthoz et al., 2002), reading stories with characters that violated social norms activated the medial prefrontal cortex as well as the neighboring lateral orbitofrontal cortex which is often activated in response to aversive emotional expressions such as anger. Notably, this neural activity was significantly higher when characters intentionally violated social norms (“Joanna has a bite of the first course, but does not like it and spits the food back into her plate”) than when they unintentionally violated them (“Joanna has a bite of the first course, chokes, and spits out the food while she is coughing”).

These studies reveal that stories engage people to understand intents and motivations in order to get inside the characters’ heads. In contrast, people are less engaged by straightforward, factual information that is not character-driven. If the storyreceiver is simply told that the tulips are dead or that a woman accidentally choked, the brain is less engaged. But if the story provides just the right amount of ambiguity or emotional inconsistency, people cannot resist sleuthing to reconcile the discrepancies and understand the character as a cohesive person with knowable mental processes.

Self-agency. Beliefs and preferences provide clues to a character’s motivations and future actions. Being able to understand that a character holds a false belief or violates a social norm provides more intelligence about their self-agency. The temporoparietal junction is engaged, with some cultural differences, to understand this self-agency. An fMRI study (Jenkins & Mitchell, 2010) revealed that the right temporoparietal junction is especially sensitive to the distinction between characters’ beliefs (“she thinks her flowers have died”) and preferences (“she always sits in the back of the class”), regardless of the ambiguity of story inferences. Furthermore, another fMRI study (Gobbini & Koralek, 2007) found that realizing a character holds a false belief activates the temporoparietal junction. For instance, to truly understand a character’s self-agency, the storyreceiver would need to realize that a character did not know their drink was poisoned when they gulped it down. The social norms fMRI study (Berthoz et al., 2002) also found activity in the temporoparietal junction for stories with characters violating social norms as part of their self-agency.

The role of the temporoparietal junction is not language-specific, reinforcing that people understand stories from a character-oriented, mentalistic perspective. In an fMRI study (Yuan et al, 2018), participants received headline-like short stories (“surgeon finds scissors inside of patient”). fMRI revealed that the temporoparietal junction was activated whether participants represented the character’s actions with words, gestures, or drawings.

However, the role of the temporoparietal junction, and thus the role of characters’ self-agency in the story framework, is culture-specific. An fMRI study (Kobayashi et al., 2006) with American and Japanese participants found that only American storyreceivers had significant activity in the temporoparietal junction. This suggests that the American culture of raising children on individualist self-agency activates the temporoparietal junction, while the Japanese culture of raising children on group-agency does not.

Social Interaction. In order to understand how characters socially engage, storyreceivers must also understand the mental states of multiple characters who interact with each other. This takes Theory of Mind a step further: storyreceivers mentalize characters who mentalize other characters. A network of brain areas helps the storyreceiver understand and infer the thoughts, feelings, motivations, and social relations of characters such as decoding irony and sarcasm by understanding the characters’ perspectives (AbdulSabur et al., 2014). These brain areas include the precuneus, invoked in self-consciousness and self-awareness; the inferior parietal lobule, interpreting sensory information including facial emotions; and the parahippocampal gyri that play a role in spatial memory and navigation (Rajimehr & Tootell, 2008). The combined fMRI and PET study (AbdulSabur et al., 2014) revealed that the precuneus, inferior parietal lobules, and parahippocampal gyri form a left-laterally activated network, along with the medial prefrontal cortex, during storytelling and storyreceiving. This network of activation illustrates that social relationships are an important element of Theory of Mind. On story, the brain must understand the mental states of multiple characters in order to understand how they will engage socially.

Actions

The brain treats stories as reality because characters trigger similar neural activity as if the storyreceiver was performing the action or feeling the same way. Hence, during story comprehension and production, the motor cortex is activated to mentally simulate the actions depicted in the story (Mar, 2004). As the frontal lobe is goal-centric, motor neural networks are more highly related to goals of action than to specific muscle movements.

The primary motor cortex, covering the posterior frontal lobe, collects input from all cortical areas involved in motor control. An fMRI study (Boulenger et al., 2009) revealed that story language activates specific primary motor cortex areas related to characters performing specific actions. Notably, sentences such as “John grasped the object” more strongly activated the lateral motor cortex, which is more involved in arm movement; sentences such as “Pablo kicked the ball” more strongly activated the dorsal motor cortex, which is more involved in leg movement.

Secondary motor areas help plan and control movement. These areas include the supplementary motor area and the premotor cortex. The supplementary motor area has strong connections to the medial frontal cortex for internally-guided goals based on personal experiences. During a story, the pre-supplementary motor area reconciles conflicting or inconsistent information about characters’ actions (AbdulSabur et al., 2014). It helps execute working memory, monitor behavioral sequences, and represent space and time in the brain. Meanwhile, the premotor cortex has strong connections to the parietal lobe for sensory, externally-guided action. When characters interact with objects, such as picking them up or putting them down, premotor cortex regions involved in the human grasping circuit are activated (Speer et al., 2012). Notably, the premotor cortex is more active when comprehending stories rather than disconnected words or sentences (AbdulSabur et al., 2014).

Extending from the premotor cortex out into the parietal and temporal lobes, mirror neurons fire in the brain when watching someone else perform an action, as if one was performing the action themselves. Mirror neurons also fire when experiencing stories through a character, allowing the storyreceiver to create powerful story simulations in their minds (Gottschall, 2012). This helps the storyreceiver viscerally feel the character’s experience, as if it was happening to them, in order to understand the goals of their actions.

Character-driven Plot

The story framework requires a character-driven plot, focusing on how a character changes, learns, and grows because of a conflict impeding their goal. Stories need a goal-driven character with some inner driving force because characters provide a viewpoint for the storyreceiver to understand the story information and events, making the plot relevant, meaningful, and memorable. People automatically assess characters’ decisions and actions to determine the characters’ intentions (Haven, 2007, p. 41). Furthermore, the plot’s conflicts must be specific to the protagonist’s goal. Stories can only captivate if a character reaches a challenge with no predefined solution. Cognitively, the story becomes an emotional experience by engaging characters while they face these challenges (Oatley, 2013). Because the brain assumes characters act rationally to achieve their goals, their actions in the plot must have intentions based on their goals and motivations. Storyreceivers focus on the characters’ goal-oriented actions due to the survival-based evolutionary role of story in acquiring insight to another person’s upcoming behavior. If a goal is achieved, the storyreceiver stops paying attention to those actions and focuses on a new goal state; if a goal is paused, the storyreceiver redirects towards a more active, relevant goal (Boyd, 2010, p. 158).

Specific brain areas track and respond to these changes in characters’ goals or characters altogether. An fMRI study (Speer et al., 2012) discovered that changes in characters or their goals bilaterally activate the posterior superior temporal cortex. This brain area is known for being activated more by goal-directed, intentional actions than aimless actions. Additionally, the lateral prefrontal cortex is activated by changes in characters’ goals while the medial prefrontal cortex is activated by changes in characters altogether. This illustrates that on story, the brain keeps careful track of how people are acting and what they are working towards, reinforcing the evolutionary advantage of understanding what others want and how they will act to get it.

Mental Representational Model

People build character-centric multimodal mental representations of stories. Built from sensory input (Gopnik et al., 1999), mental representations interpret and incorporate the onslaught of story events. Schemas help assimilate the story’s events into this mental construction that considers the storyreceivers’ previous knowledge, attitudes, and feelings to make the story comprehensible (Oatley, 2013). fMRI studies reveal that storyreceivers’ situational models are similar to the brain processes to recall previous situations or imagine hypothetical situations (Speer et al., 2012).

Story modality doesn’t restrict how the inner representation is formed. Multimodal, though perhaps visual-heavy, representations and simulations of a story are formed to understand its perceptual, motor, and affective content (Barsalou, 2008). During story processing, the posterior cingulate cortex retrieves elaborative information including visuospatial imagery, episodic retrieval, and emotional modulation of memory processes (Mar, 2004). PET scans reveal that stories activate the posterior cingulate cortex to help encode relevant story material in episodic memory, likely in a mostly visual modality (Fletcher et al., 1995). This helps build a cohesive narrative that can incorporate future events into the story structure. The elaborative role of the posterior cingulate cortex supports autonoetic awareness, which is one’s ability to place themselves in the past, future, or fictional situations, and examine their own thoughts (Mar, 2004).

A PET study (Maguire et al., 1999) found posterior cingulate activation when participants associated incoming information with prior knowledge to create a coherent narrative representation. The study provided the overall gist of the story upfront, giving participants prior knowledge, and then presented usual stories (“Long ago a musician was astounded when his four-year-old son wrote out a concerto for harpsichord…”) or unusual stories (“If the balloons popped the sound would not be able to carry since everything would be away from the correct floor…Since the whole operation depends on a steady flow of electricity…”). Even the unusual stories activated the posterior cingulate cortex, highlighting the importance of prior knowledge in building comprehensible stories.

Story Sequencing

As each story beat leads to the next, the storyteller must draw cause and effect between story actions. When a story sparks curiosity, the brain’s capacity for learning increases. Curiosity in turn sparks suspense, which releases dopamine, a pleasurable brain chemical that helps with movement, attention, learning, and emotions (Gruber et al., 2014). Suspense also releases cortisol, a stress hormone that commands one to pay attention (Zak, 2015).

Because people assume all story events are connected, the brain will mentally invent anything to ensure the story makes sense via cause-and effect and temporal sequencing (Haven, 2007). If any story content turns out to be extraneous, the brain either stops firing dopamine, signaling to the storyreceiver their anticipation was for naught (Lehrer, 2009), or jumps to conclusions about the story based on available information. Consequently, as Golden (2017) advises, a good story “relies largely on how adeptly you zoom in and zoom out across time. So strike the right balance, soaring high for perspective, and going lower so we can see the ground” (p. 104). Neuroimaging studies reveal that several brain areas work together to construct a cohesive, coherent story by selecting and sequencing story information.

Temporal Lobe. The temporal lobe serves as an intermediary between sensory input and the representational model (Ferstl et al., 2014). The left temporal gyrus and the temporal poles support language comprehension including concatenating sentences. Notably, these brain areas are not involved in simple semantic processes such as sentence processing (Mar, 2004). Several studies reinforce their role in understanding story language content. A PET study (Fletcher et al., 1995) found that compared to random sentences, stories activate the temporal poles bilaterally in order to link story propositions together to build a cohesive, visceral representation of the story. Additionally, stories activate the left superior temporal gyrus, which makes language cohesive across sentence boundaries. A PET study (Maguire et al., 1999) found the left temporal pole is activated to help create a narrative by concatenating sentences. In another PET study (Mazoyer et al., 1993), French participants listened to meaningful or distorted stories with nonsense, unrelated, or non-French words. PET scans revealed that processing meaningful stories activated the left middle temporal gyrus and temporal poles. In the combined fMRI and PET study by AbdulSabur et al. (2014), both storytelling and storyreceiving activated the temporal poles. This demonstrates that temporal pole activation is linked to processing more complex discourse-level semantic knowledge and concatenating sentences during storytelling or storyreceiving.

Frontoparietal Network. The parietal lobe contains part of the frontoparietal attention system that is linked to motor intention and movement goals. As already established, the premotor cortex has strong ties to the parietal lobe for sensory, externally-guided actions. In an fMRI study (Ferstl et al., 2014), the dorsal part of the anterior precuneus and intraparietal sulci helped build a cohesive story chronology, in partnership with the goal-oriented, mentalistic middle frontal gyri. This finding reflects the frontoparietal networks’ role in working memory and attentional processes. The dorsal precuneus is activated by switches from local to global stimuli while the intraparietal sulci and middle frontal gyri facilitate verbal working memory tasks during articulatory suppression. This is relevant because storyreceivers must hold story content in working memory while continuing to comprehend the next story event being presented.

Frontal Lobe. The frontal lobe supports construction of global coherence for a story. It is involved in short-term memory, problem solving, creativity, and judgement (Żurawicki, 2010). An fMRI study by Stephens and colleagues (2010) revealed synchronized prefrontal brain activity between storyteller and storyreceiver. Synchronized areas included the dorsolateral prefrontal cortex, orbitofrontal cortex, and medial prefrontal cortex. Notably, prefrontal activity in the storyreceiver preceded the storyteller’s activity, suggesting that the prefrontal cortex anticipates upcoming story input that will need processing.

The prefrontal cortex represents structured, goal-oriented, sequential events (Grafman, 2002). The medial prefrontal cortex and lateral prefrontal cortex select and sequence story information to produce a representation in working memory (Mar, 2004). In particular, the dorsolateral prefrontal cortex plays a key role in ordering story events over a period of time. Meanwhile, the medial prefrontal cortex retrieves information about when, where, and how events occurred and extracts themes for global coherence (AbdulSabur et al., 2014). These working memory processes are bolstered by two frontal areas that help modulate attentional focus to the story: the orbitofrontal cortex and anterior cingulate cortex (Mar, 2004). The orbitofrontal cortex regulates planned behavior, anticipating reward and punishment (Żurawicki, 2010). A PET study (Maguire et al., 1999) revealed that ventromedial orbitofrontal cortex activation increases as story comprehension grows. The anterior cingulate cortex, wrapped above the corpus callosum that connects the two brain hemispheres, works with the limbic system for emotional processing, learning, and memory (Żurawicki, 2010).

Moral of the Story

At some point, the story descends to its darkest point, where the protagonist is most threatened or helpless to reach their goal. Emerging from the darkest point of the plot, the story framework does not require the character to reach their goal. Story closure could be the realization that the goal is unreachable, or the events come to a natural end (Ohler, 2013). More important is that the protagonist’s inner self is changed in some way by the steps they took to try to achieve their goal (Cron, 2012). Change is the catalyst that makes stories worthwhile and their key message more memorable (Golden, 2017). Two brain areas lead this effort. In the combined fMRI and PET study (AbdulSabur et al., 2014), the posterior middle temporal gyrus partnered with the inferior frontal gyrus for higher-level semantic retrieval and integration in order to understand a story’s loosely-associated information such as themes and morals. Another PET study (Nichelli et al., 1995) using Aesop’s fables also illuminated that storyreceivers use the right inferior frontal gyrus, in conjunction with the right middle temporal gyrus, to comprehend the moral of the story. Notably, to grasp the theme or moral, these two brain areas must encode a compilation of individual story events as they relate to the characters of the story.

Persuasive Power

The story framework helps explain the immense persuasive power of stories on people. Stories change affective and cognitive responses, beliefs, attitudes, and intentions (Van Laer et al., 2014). Yet, people are so hardwired for story that they are often unaware of their effects. As neuromarketers Morin and Renvoise explain, stories “have the power to reshape the beliefs and behaviors of the listeners because they fool the story receiver’s primal brain in believing the story is real” (2018, p. 184). Transported storyreceivers are more likely to accept the story as authentic and less likely to identify any falsehoods in the story (Green & Brock, 2013). This illustrates what neuromarketers know best: people’s choices are framed without their knowledge, their emotions restrict rationality, and decisions are instantaneous, not deliberate (Ramsøy, 2015). Because stories are mostly automatic and subconscious, they have a stealth influence on people.

Story persuasion leads to belief, attitude, and intention changes that last longer and are more shielded from counter-influence than rhetoric, argumentation, or other forms of discourse (Green & Brock, 2013). Much research is now directed towards the cultural, economic and ethical implications of using stories to influence consumer decision-making (Van Laer et al., 2014). Consumer research reveals that story characters can increase storyreceievers’ intentions to eat healthier (Slater et al., 2003), start smoking (Dal Cin et al., 2007), purchase an MP3 player (Van den Hende et al., 2007) or digital camera (Schlosser, 2003), or use sunscreen. Stories also impact beliefs over time. In a study (Appel & Richter, 2007), participants who read a fictional story with false assertions about real-world topics. Storyreceivers’ belief of the false story elements was even stronger when measured two weeks later, compared to immediately after the storytelling occurred.

Stories allow for empathy because their characters simulate unknown people in the ordinary world (Oatley, 2013). Because the storyreceiver experiences the character as a real person who seems to be trusted, safe, and familiar, the hypothalamus releases oxytocin (Zak, 2015). Oxytocin is a chemical that improves empathy, healing, and blood pressure while motivating reciprocation and pro-social behaviors. In a study (Barraza & Zak, 2009), participants watched one of two videos. In a story-based video set at a hospital, a father speaks about Ben, his bald two-year-old son with terminal brain cancer who is playing the background. In a non-story-based video, Ben and his father spend a day at the zoo, but his illness is not mentioned. As measured by blood samples, the story-based video set in the hospital induced cortisol and oxytocin, increasing empathy for the characters, and prompted the storyreceiver to engage donate to charity. The non-story-based video did not have any of the same effects. In a follow-up study (Barraza et al., 2015), viewers’ attention was measured using heart rate via electrocardiogram (ECG) and sweat via an electrodermal sensor on the fingers. In addition, ECG captured viewers’ emotional resonance by measuring activity in the vagus nerve, which is dense in oxytocin receptors. Early in the video, vagal activity increased as the viewer was acquainted with Ben and his father; attention peaked at the video’s climax when the father reveals Ben is dying. Viewers who attended to the video and created an emotional connection were almost certain to donate to charity; in fact, brain activity predicted with 82% accuracy if a storyreceiver would donate.

A different study (Lin et al., 2013) measured viewer responses to public service announcements (PSAs) presented as video stories. In the first experiment, viewers who received synthetic oxytocin through their noses donated 56% more money to charity than viewers who received a placebo. Lin et al. (2013) concluded that oxytocin increases post-narrative prosocial behavior because it increases empathy for the characters in the PSAs. In the second experiment, viewers who paid attention to the PSAs had increased levels of adrenocorticotropin (an arousal hormone) and oxytocin in their blood, which led them to donate 261% more to charity than participants who did not pay attention to the public service announcements. Amazingly, although viewers knew the characters in the PSAs were actors, and donations could not solve their specific fictional problems, the emotional connection still triggered prosocial behavior.

Conclusion

Today, stories permeate movies, television, social media, websites, blogs, data visualizations, podcasts, video games, virtual reality, augmented reality, advertisements, and more. While digital-age media technologies afford storytelling that is created faster and travels further than ever before, the human brain remains the same. Hence, technology is merely an amplifier for story, not a fixer of story (Ohler, 2013, p. 6). Media storytellers must still consider the brain’s time-honored and evolutionarily-guided rules for story. The neuroscientific theoretical framework presented here can help digital-age storytellers understand and predict how their media stories will engage and affect audiences. The story framework includes emotionally-engaging characters with knowable mental processes who act with intent and motive. With this cast of goal-driven characters, story framework requires character-driven plot that focuses on how someone changes, learns, and grows because of a conflict impeding their goal. From the character-driven plot, people build a multimodal representation of the story to interpret and incorporate the onslaught of story events, building empathy and uncovering themes and morals that can change attitudes or beliefs. Ultimately, this framework helps storytellers craft a compelling story in the digital age that engages people, provides a round-trip ticket into the storyworld, and leaves a lasting impression.

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