Emotions, fast and slow: processing of emotion words is affected by individual differences in need for affect and narrative absorption.

Emotional words have consistently been shown to be processed differently than neutral words. However, few studies have examined individual variability in emotion word processing with longer, ecologically valid stimuli (beyond isolated words, sentences, or paragraphs). In the current study, we re-analysed eye-tracking data collected during story reading to reveal how individual differences in need for affect and narrative absorption impact the speed of emotion word reading. Word emotionality was indexed by affective-aesthetic potentials (AAP) calculated by a sentiment analysis tool. We found that individuals with higher levels of need for affect and narrative absorption read positive words more slowly. On the other hand, these individual differences did not influence the reading time of more negative words, suggesting that high need for affect and narrative absorption are characterised by a positivity bias only. In general, unlike most previous studies using more isolated emotion word stimuli, we observed a quadratic (U-shaped) effect of word emotionality on reading speed, such that both positive and negative words were processed more slowly than neutral words. Taken together, this study emphasises the importance of taking into account individual differences and task context when studying emotion word processing.

The Storytelling Brain: How Neuroscience Stories Help Bridge the Gap between Research and Society.

Active communication between researchers and society is necessary for the scientific community's involvement in developing science-based policies. This need is recognized by governmental and funding agencies that compel scientists to increase their public engagement and disseminate research findings in an accessible fashion. Storytelling techniques can help convey science by engaging people's imagination and emotions. Yet, many researchers are uncertain about how to approach scientific storytelling, or feel they lack the tools to undertake it. Here we explore some of the techniques intrinsic to crafting scientific narratives, as well as the reasons why scientific storytelling may be an optimal way of communicating research to nonspecialists. We also point out current communication gaps between science and society, particularly in the context of neurodiverse audiences and those that include neurological and psychiatric patients. Present shortcomings may turn into areas of synergy with the potential to link neuroscience education, research, and advocacy.

Frequency-specific brain dynamics related to prediction during language comprehension.

The brain's remarkable capacity to process spoken language virtually in real time requires fast and efficient information processing machinery. In this study, we investigated how frequency-specific brain dynamics relate to models of probabilistic language prediction during auditory narrative comprehension. We recorded MEG activity while participants were listening to auditory stories in Dutch. Using trigram statistical language models, we estimated for every word in a story its conditional probability of occurrence. On the basis of word probabilities, we computed how unexpected the current word is given its context (word perplexity) and how (un)predictable the current linguistic context is (word entropy). We then evaluated whether source-reconstructed MEG oscillations at different frequency bands are modulated as a function of these language processing metrics. We show that theta-band source dynamics are increased in high relative to low entropy states, likely reflecting lexical computations. Beta-band dynamics are increased in situations of low word entropy and perplexity possibly reflecting maintenance of ongoing cognitive context. These findings lend support to the idea that the brain engages in the active generation and evaluation of predicted language based on the statistical properties of the input signal.

On the other hand: including left-handers in cognitive neuroscience and neurogenetics.

Left-handers are often excluded from study cohorts in neuroscience and neurogenetics in order to reduce variance in the data. However, recent investigations have shown that the inclusion or targeted recruitment of left-handers can be informative in studies on a range of topics, such as cerebral lateralization and the genetic underpinning of asymmetrical brain development. Left-handed individuals represent a substantial portion of the human population and therefore left-handedness falls within the normal range of human diversity; thus, it is important to account for this variation in our understanding of brain functioning. We call for neuroscientists and neurogeneticists to recognize the potential of studying this often-discarded group of research subjects.

Prediction During Natural Language Comprehension.

The notion of prediction is studied in cognitive neuroscience with increasing intensity. We investigated the neural basis of 2 distinct aspects of word prediction, derived from information theory, during story comprehension. We assessed the effect of entropy of next-word probability distributions as well as surprisal A computational model determined entropy and surprisal for each word in 3 literary stories. Twenty-four healthy participants listened to the same 3 stories while their brain activation was measured using fMRI. Reversed speech fragments were presented as a control condition. Brain areas sensitive to entropy were left ventral premotor cortex, left middle frontal gyrus, right inferior frontal gyrus, left inferior parietal lobule, and left supplementary motor area. Areas sensitive to surprisal were left inferior temporal sulcus ("visual word form area"), bilateral superior temporal gyrus, right amygdala, bilateral anterior temporal poles, and right inferior frontal sulcus. We conclude that prediction during language comprehension can occur at several levels of processing, including at the level of word form. Our study exemplifies the power of combining computational linguistics with cognitive neuroscience, and additionally underlines the feasibility of studying continuous spoken language materials with fMRI.

Feel between the lines: implied emotion in sentence comprehension.

This study investigated the brain regions for the comprehension of implied emotion in sentences. Participants read negative sentences without negative words, for example, "The boy fell asleep and never woke up again," and their neutral counterparts "The boy stood up and grabbed his bag." This kind of negative sentence allows us to examine implied emotion derived at the sentence level, without associative emotion coming from word retrieval. We found that implied emotion in sentences, relative to neutral sentences, led to activation in some emotion-related areas, including the medial prefrontal cortex, the amygdala, and the insula, as well as certain language-related areas, including the inferior frontal gyrus, which has been implicated in combinatorial processing. These results suggest that the emotional network involved in implied emotion is intricately related to the network for combinatorial processing in language, supporting the view that sentence meaning is more than simply concatenating the meanings of its lexical building blocks.

Beyond the tried and true: How virtual reality, dialog setups, and a focus on multimodality can take bilingual language production research forward.

Bilinguals possess the ability of expressing themselves in more than one language, and typically do so in contextually rich and dynamic settings. Theories and models have indeed long considered context factors to affect bilingual language production in many ways. However, most experimental studies in this domain have failed to fully incorporate linguistic, social, or physical context aspects, let alone combine them in the same study. Indeed, most experimental psycholinguistic research has taken place in isolated and constrained lab settings with carefully selected words or sentences, rather than under rich and naturalistic conditions. We argue that the most influential experimental paradigms in the psycholinguistic study of bilingual language production fall short of capturing the effects of context on language processing and control presupposed by prominent models. This paper therefore aims to enrich the methodological basis for investigating context aspects in current experimental paradigms and thereby move the field of bilingual language production research forward theoretically. After considering extensions of existing paradigms proposed to address context effects, we present three far-ranging innovative proposals, focusing on virtual reality, dialog situations, and multimodality in the context of bilingual language production.

MA-EM: A neurocognitive model for understanding mixed and ambiguous emotions and morality.

Understanding emotions and moral intentions of other people is integral to being human. Humanities scholars have long recognized the complex and ambiguous nature of emotions and morality. People are rarely 'just' happy, or sad. Neither are they 'just' good or bad people. Despite this, most knowledge about the psychological and neural basis of emotions and moral understanding comes from experiments investigating unidimensional and non-ambiguous emotions and morality. The goal of this paper is twofold. First I want to point out why mixed and ambiguous emotions and morality are a promising research topic for cognitive neuroscientists. Observing or experiencing mixed or ambiguous emotions and morality tends to have a strong impact on humans. This impact is clearly visible in narratives and fiction, and I will argue that narratives make an excellent stimulus to study the effects of emotional and moral ambiguity. Second, I will sketch a model to help guide research in this promising corner of human cognition.

Different kinds of simulation during literary reading: Insights from a combined fMRI and eye-tracking study.

Mental simulation is an important aspect of narrative reading. In a previous study, we found that gaze durations are differentially impacted by different kinds of mental simulation. Motor simulation, perceptual simulation, and mentalizing as elicited by literary short stories influenced eye movements in distinguishable ways (Mak & Willems, 2019). In the current study, we investigated the existence of a common neural locus for these different kinds of simulation. We additionally investigated whether individual differences during reading, as indexed by the eye movements, are reflected in domain-specific activations in the brain. We found a variety of brain areas activated by simulation-eliciting content, both modality-specific brain areas and a general simulation area. Individual variation in percent signal change in activated areas was related to measures of story appreciation as well as personal characteristics (i.e., transportability, perspective taking). Taken together, these findings suggest that mental simulation is supported by both domain-specific processes grounded in previous experiences, and by the neural mechanisms that underlie higher-order language processing (e.g., situation model building, event indexing, integration).

Reading about minds: The social-cognitive potential of narratives.

It is often argued that narratives improve social cognition, either by appealing to social-cognitive abilities as we engage with the story world and its characters, or by conveying social knowledge. Empirical studies have found support for both a correlational and a causal link between exposure to (literary, fictional) narratives and social cognition. However, a series of failed replications has cast doubt on the robustness of these claims. Here, we review the existing empirical literature and identify open questions and challenges. An important conclusion of the review is that previous research has given too little consideration to the diversity of narratives, readers, and social-cognitive processes involved in the social-cognitive potential of narratives. We therefore establish a research agenda, proposing that future research should focus on (1) the specific text characteristics that drive the social-cognitive potential of narratives, (2) the individual differences between readers with respect to their sensitivity to this potential, and (3) the various aspects of social cognition that are potentially affected by reading narratives. Our recommendations can guide the design of future studies that will help us understand how, for whom, and in what respect exposure to narratives can advantage social cognition.

Different routes to liking: how readers arrive at narrative evaluations.

When two people read the same story, they might both end up liking it very much. However, this does not necessarily mean that their reasons for liking it were identical. We therefore ask what factors contribute to "liking" a story, and-most importantly-how people vary in this respect. We found that readers like stories because they find them interesting, amusing, suspenseful and/or beautiful. However, the degree to which these components of appreciation were related to how much readers liked stories differed between individuals. Interestingly, the individual slopes of the relationships between many of the components and liking were (positively or negatively) correlated. This indicated, for instance, that individuals displaying a relatively strong relationship between interest and liking, generally display a relatively weak relationship between sadness and liking. The individual differences in the strengths of the relationships between the components and liking were not related to individual differences in expertize, a characteristic strongly associated with aesthetic appreciation of visual art. Our work illustrates that it is important to take into consideration the fact that individuals differ in how they arrive at their evaluation of literary stories, and that it is possible to quantify these differences in empirical experiments. Our work suggests that future research should be careful about "overfitting" theories of aesthetic appreciation to an "idealized reader," but rather take into consideration variations across individuals in the reason for liking a particular story.

Complementary systems for understanding action intentions.

How humans understand the intention of others' actions remains controversial. Some authors have suggested that intentions are recognized by means of a motor simulation of the observed action with the mirror-neuron system [1-3]. Others emphasize that intention recognition is an inferential process, often called "mentalizing" or employing a "theory of mind," which activates areas well outside the motor system [4-6]. Here, we assessed the contribution of brain regions involved in motor simulation and mentalizing for understanding action intentions via functional brain imaging. Results show that the inferior frontal gyrus (part of the mirror-neuron system) processes the intentionality of an observed action on the basis of the visual properties of the action, irrespective of whether the subject paid attention to the intention or not. Conversely, brain areas that are part of a "mentalizing" network become active when subjects reflect about the intentionality of an observed action, but they are largely insensitive to the visual properties of the observed action. This supports the hypothesis that motor simulation and mentalizing have distinct but complementary functions for the recognition of others' intentions.

Altered intrinsic functional connectivity of anterior and posterior insula regions in high-functioning participants with autism spectrum disorder.

Impaired understanding of others' sensations and emotions as well as abnormal experience of their own emotions and sensations is frequently reported in individuals with Autism Spectrum Disorder (ASD). It is hypothesized that these abnormalities are based on altered connectivity within "shared" neural networks involved in emotional awareness of self and others. The insula is considered a central brain region in a network underlying these functions, being located at the transition of information about bodily arousal and the physiological state of the body to subjective feelings. The present study investigated the intrinsic functional connectivity properties of the insula in 14 high-functioning participants with ASD (HF-ASD) and 15 typically developing (TD) participants in the age range between 12 and 20 years by means of "resting state" or "nontask" functional magnetic resonance imaging. Essentially, a distinction was made between anterior and posterior regions of the insular cortex. The results show a reduced functional connectivity in the HF-ASD group, compared with the TD group, between anterior as well as posterior insula and specific brain regions involved in emotional and sensory processing. It is suggested that functional abnormalities in a network involved in emotional and interoceptive awareness might be at the basis of altered emotional experiences and impaired social abilities in ASD, and that these abnormalities are partly based on the intrinsic functional connectivity properties of such a network.

A functional role for the motor system in language understanding: evidence from theta-burst transcranial magnetic stimulation.

Does language comprehension depend, in part, on neural systems for action? In previous studies, motor areas of the brain were activated when people read or listened to action verbs, but it remains unclear whether such activation is functionally relevant for comprehension. In the experiments reported here, we used off-line theta-burst transcranial magnetic stimulation to investigate whether a causal relationship exists between activity in premotor cortex and action-language understanding. Right-handed participants completed a lexical decision task, in which they read verbs describing manual actions typically performed with the dominant hand (e.g., "to throw," "to write") and verbs describing nonmanual actions (e.g., "to earn," "to wander"). Responses to manual-action verbs (but not to nonmanual-action verbs) were faster after stimulation of the hand area in left premotor cortex than after stimulation of the hand area in right premotor cortex. These results suggest that premotor cortex has a functional role in action-language understanding.

Cerebral lateralization of face-selective and body-selective visual areas depends on handedness.

The left-hemisphere dominance for language is a core example of the functional specialization of the cerebral hemispheres. The degree of left-hemisphere dominance for language depends on hand preference: Whereas the majority of right-handers show left-hemispheric language lateralization, this number is reduced in left-handers. Here, we assessed whether handedness analogously has an influence upon lateralization in the visual system. Using functional magnetic resonance imaging, we localized 4 more or less specialized extrastriate areas in left- and right-handers, namely fusiform face area (FFA), extrastriate body area (EBA), fusiform body area (FBA), and human motion area (human middle temporal [hMT]). We found that lateralization of FFA and EBA depends on handedness: These areas were right lateralized in right-handers but not in left-handers. A similar tendency was observed in FBA but not in hMT. We conclude that the relationship between handedness and hemispheric lateralization extends to functionally lateralized parts of visual cortex, indicating a general coupling between cerebral lateralization and handedness. Our findings indicate that hemispheric specialization is not fixed but can vary considerably across individuals even in areas engaged relatively early in the visual system.

How context changes the neural basis of perception and language.

Cognitive processes-from basic sensory analysis to language understanding-are typically contextualized. While the importance of considering context for understanding cognition has long been recognized in psychology and philosophy, it has not yet had much impact on cognitive neuroscience research, where cognition is often studied in decontextualized paradigms. Here, we present examples of recent studies showing that context changes the neural basis of diverse cognitive processes, including perception, attention, memory, and language. Within the domains of perception and language, we review neuroimaging results showing that context interacts with stimulus processing, changes activity in classical perception and language regions, and recruits additional brain regions that contribute crucially to naturalistic perception and language. We discuss how contextualized cognitive neuroscience will allow for discovering new principles of the mind and brain.

Reading Minds, Reading Stories: Social-Cognitive Abilities Affect the Linguistic Processing of Narrative Viewpoint.

Although various studies have shown that narrative reading draws on social-cognitive abilities, not much is known about the precise aspects of narrative processing that engage these abilities. We hypothesized that the linguistic processing of narrative viewpoint-expressed by elements that provide access to the inner world of characters-might play an important role in engaging social-cognitive abilities. Using eye tracking, we studied the effect of lexical markers of perceptual, cognitive, and emotional viewpoint on eye movements during reading of a 5,000-word narrative. Next, we investigated how this relationship was modulated by individual differences in social-cognitive abilities. Our results show diverging patterns of eye movements for perceptual viewpoint markers on the one hand, and cognitive and emotional viewpoint markers on the other. Whereas the former are processed relatively fast compared to non-viewpoint markers, the latter are processed relatively slow. Moreover, we found that social-cognitive abilities impacted the processing of words in general, and of perceptual and cognitive viewpoint markers in particular, such that both perspective-taking abilities and self-reported perspective-taking traits facilitated the processing of these markers. All in all, our study extends earlier findings that social cognition is of importance for story reading, showing that individual differences in social-cognitive abilities are related to the linguistic processing of narrative viewpoint.

Distinguishing Syntactic Operations in the Brain: Dependency and Phrase-Structure Parsing.

Finding the structure of a sentence-the way its words hold together to convey meaning-is a fundamental step in language comprehension. Several brain regions, including the left inferior frontal gyrus, the left posterior superior temporal gyrus, and the left anterior temporal pole, are supposed to support this operation. The exact role of these areas is nonetheless still debated. In this paper we investigate the hypothesis that different brain regions could be sensitive to different kinds of syntactic computations. We compare the fit of phrase-structure and dependency structure descriptors to activity in brain areas using fMRI. Our results show a division between areas with regard to the type of structure computed, with the left anterior temporal pole and left inferior frontal gyrus favouring dependency structures and left posterior superior temporal gyrus favouring phrase structures.

Neural dissociations between action verb understanding and motor imagery.

According to embodied theories of language, people understand a verb like throw, at least in part, by mentally simulating throwing. This implicit simulation is often assumed to be similar or identical to motor imagery. Here we used fMRI to test whether implicit simulations of actions during language understanding involve the same cortical motor regions as explicit motor imagery. Healthy participants were presented with verbs related to hand actions (e.g., to throw) and nonmanual actions (e.g., to kneel). They either read these verbs (lexical decision task) or actively imagined performing the actions named by the verbs (imagery task). Primary motor cortex showed effector-specific activation during imagery, but not during lexical decision. Parts of premotor cortex distinguished manual from nonmanual actions during both lexical decision and imagery, but there was no overlap or correlation between regions activated during the two tasks. These dissociations suggest that implicit simulation and explicit imagery cued by action verbs may involve different types of motor representations and that the construct of "mental simulation" should be distinguished from "mental imagery" in embodied theories of language.

The function of words: distinct neural correlates for words denoting differently manipulable objects.

Recent research indicates that language processing relies on brain areas dedicated to perception and action. For example, processing words denoting manipulable objects has been shown to activate a fronto-parietal network involved in actual tool use. This is suggested to reflect the knowledge the subject has about how objects are moved and used. However, information about how to use an object may be much more central to the conceptual representation of an object than information about how to move an object. Therefore, there may be much more fine-grained distinctions between objects on the neural level, especially related to the usability of manipulable objects. In the current study, we investigated whether a distinction can be made between words denoting (1) objects that can be picked up to move (e.g., volumetrically manipulable objects: bookend, clock) and (2) objects that must be picked up to use (e.g., functionally manipulable objects: cup, pen). The results show that functionally manipulable words elicit greater levels of activation in the fronto-parietal sensorimotor areas than volumetrically manipulable words. This suggests that indeed a distinction can be made between different types of manipulable objects. Specifically, how an object is used functionally rather than whether an object can be displaced with the hand is reflected in semantic representations in the brain.