The neurobiological basis of affect is consistent with psychological construction theory and shares a common neural basis across emotional categories.

Affective experience colours everyday perception and cognition, yet its fundamental and neurobiological basis is poorly understood. The current debate essentially centers around the communalities and specificities across individuals, events, and emotional categories like anger, sadness, and happiness. Using fMRI during the experience of these emotions, we critically compare the two dominant conflicting theories on human affect. Basic emotion theory posits emotions as discrete universal entities generated by dedicated emotion category-specific neural circuits, while psychological construction theory claims emotional events as unique, idiosyncratic, and constructed by psychological primitives like core affect and conceptualization, which underlie each emotional event and operate in a predictive framework. Based on the findings of 8 a priori-defined model-specific prediction tests on the neural response amplitudes and patterns, we conclude that the neurobiological basis of affect is primarily characterized by idiosyncratic mechanisms and a common neural basis shared across emotion categories, consistent with psychological construction theory. The findings provide further insight into the organizational principles of the neural basis of affect and brain function in general. Future studies in clinical populations with affective symptoms may reveal the corresponding underlying neural changes from a psychological construction perspective.

Maternal dopamine encodes affective signals of human infants.

Mothers are highly responsive to their offspring. In non-human mammals, mothers secrete dopamine in the nucleus accumbens (NAcc) in response to their pups. Yet, it is still unknown which aspect of the offspring behavior elicits dopaminergic responses in mothers. Here, we tested whether infants' affective signals elicit dopaminergic responses in the NAcc of human mothers. First, we conducted a behavioral analysis on videos of infants' free play and quantified the affective signals infants spontaneously communicated. Then, we presented the same videos to mothers during a magnetic resonance-positron emission tomography scan. We traced the binding of [11C]raclopride to free D2/3-type receptors to assess maternal dopaminergic responses during the infant videos. When mothers observed videos with many infant signals during the scan, they had less [11C]raclopride binding in the right NAcc. Less [11C]raclopride binding indicates that less D2/3 receptors were free, possibly due to increased endogenous dopamine responses to infants' affective signals. We conclude that NAcc D2/3 receptors are involved in maternal responsiveness to affective signals of human infants. D2/3 receptors have been associated with maternal responsiveness in nonhuman animals. This evidence supports a similar mechanism in humans and specifies infant-behaviors that activate the maternal dopaminergic system, with implications for social neuroscience, development and psychopathology.

A human colliculus-pulvinar-amygdala pathway encodes negative emotion.

Animals must rapidly respond to threats to survive. In rodents, threat-related signals are processed through a subcortical pathway from the superior colliculus to the amygdala, a putative "low road" to affective behavior. This pathway has not been well characterized in humans. We developed a novel pathway identification framework that uses pattern recognition to identify connected neural populations and optimize measurement of inter-region connectivity. We first verified that the model identifies known thalamocortical pathways with high sensitivity and specificity in 7 T (n = 56) and 3 T (n = 48) fMRI experiments. Then we identified a human functional superior colliculus-pulvinar-amygdala pathway. Activity in this pathway encodes the intensity of normative emotional responses to negative images and sounds but not pleasant images or painful stimuli. These results provide a functional description of a human "low road" pathway selective for negative exteroceptive events and demonstrate a promising method for characterizing human functional brain pathways.

Essentialist Biases in Reasoning About Emotions.

A large literature debates whether emotions are universal and innate. Here, we ask whether reasoning about such matters is shaped by intuitive Essentialist biases that link innateness to the material body. To gauge the perception of innateness, we asked laypeople to evaluate whether emotion categories will be recognized spontaneously by hunter-gatherers who have had no contact with Westerners. Experiment 1 shows that participants believe that emotions are innate and embodied (facially and internally) and these two properties correlate reliably. Experiment 2 demonstrates that the link is causal. When told that emotions are localized in specific brain areas (i.e., embodied), participants concluded that emotions are innate. Experiment 3 shows that this naïve view persists even when participants are explicitly informed that these emotions are acquired. Our results are the first to suggest that laypeople incorrectly believe that, if emotions are embodied, then they must be innate. We suggest that people's failure to grasp the workings of their psyche arises from the human psyche itself.

Atrophy in Distinct Corticolimbic Networks Subserving Socioaffective Behavior in Semantic Variant Primary Progressive Aphasia.

BACKGROUND: Although traditionally conceptualized as a language disorder, semantic variant primary progressive aphasia (svPPA) is often accompanied by significant behavioral and affective symptoms which considerably increase disease morbidity. Specifically, these neuropsychiatric symptoms are characterized by breaches in normative socioaffective function, for example, an inability to read social cues, excessive trusting of others, and decreased empathy. Our prior neuroimaging work identified 3 corticolimbic networks anchored in the amygdala, temporal pole, and frontoinsular cortex: an affiliation network, theorized to mediate social approach behavior; an aversion network, theorized to subserve the appraisal of social threat; and a perception network, theorized to mediate the detection of social cues. We hy-pothesized that degeneration of these networks could provide neuroanatomical substrates for socioaffective deficits in svPPA. METHODS: We examined hypothesized relationships between subscores on the Social Impairment Rating Scale (SIRS) and atrophy in each of these 3 networks in a group of 16 svPPA patients (using matched cognitively normal controls as a reference). RESULTS: Consistent with our predictions, the magnitude of atrophy in the affiliation network in svPPA patients correlated with the SIRS subscore of socioemotional detachment, while the magnitude of atrophy in the aversion network in svPPA patients correlated with the SIRS subscore of inappropriate trusting. We did not find the predicted association between perception network atrophy and the SIRS subscore of lack of attention to social cues. CONCLUSION: These findings highlight specific socioaffective deficits in svPPA and provide a neuroanatomical basis for these impairments by linking them to networks commonly targeted in this disorder.

Functional Involvement of Human Periaqueductal Gray and Other Midbrain Nuclei in Cognitive Control.

Recent theoretical advances have motivated the hypothesis that the periaqueductal gray (PAG) participates in behaviors that involve changes in the autonomic control of visceromotor activity, including during cognitively demanding tasks. We used ultra-high-field (7 tesla) fMRI to measure human brain activity at 1.1 mm resolution while participants completed a working memory task. Consistent with prior work, participants were less accurate and responded more slowly with increasing memory load-signs of increasing task difficulty. Whole-brain fMRI analysis revealed increased activity in multiple cortical areas with increasing working memory load, including frontal and parietal cortex, dorsal cingulate, supplementary motor area, and anterior insula. Several dopamine-rich midbrain nuclei, such as the substantia nigra and ventral tegmental area, also exhibited load-dependent increases in activation. To investigate PAG involvement during cognitive engagement, we developed an automated method for segmenting and spatially normalizing the PAG. Analyses using cross-validated linear support vector machines showed that the PAG discriminated high versus low working memory load conditions with 95% accuracy in individual subjects based on activity increases in lateral and ventrolateral PAG. Effect sizes in the PAG were comparable in magnitude to those in many of the cortical areas. These findings suggest that cognitive control is not only associated with cortical activity in the frontal and parietal lobes, but also with increased activity in the subcortical PAG and other midbrain regions involved in the regulation of autonomic nervous system function.SIGNIFICANCE STATEMENT Functional neuroimaging in humans has shown that cognitive control engages multiple corticostriatal networks and brainstem nuclei, but theoretical advances suggest that the periaqueductal gray (PAG) should also be engaged during cognitively demanding tasks. Recent advances in ultra-high-field fMRI provided an opportunity to obtain the first evidence that increased activation of intermediate and rostral portions of lateral and ventrolateral PAG columns in humans is modulated by cognitive load. These findings suggest that cognitive control is not solely mediated by activity in the cortex, but that midbrain structures important for autonomic regulation also play a crucial role in higher-order cognition.

Concepts dissolve artificial boundaries in the study of emotion and cognition, uniting body, brain, and mind.

Theories of emotion have often maintained artificial boundaries: for instance, that cognition and emotion are separable, and that an emotion concept is separable from the emotional events that comprise its category (e.g. "fear" is distinct from instances of fear). Over the past several years, research has dissolved these artificial boundaries, suggesting instead that conceptual construction is a domain-general process-a process by which the brain makes meaning of the world. The brain constructs emotion concepts, but also cognitions and perceptions, all in the service of guiding action. In this view, concepts are multimodal constructions, dynamically prepared from a set of highly variable instances. This approach obviates old questions (e.g. how does cognition regulate emotion?) but generates new ones (e.g. how does a brain learn emotion concepts?). In this paper, we review this constructionist, predictive coding account of emotion, considering its implications for health and well-being, culture and development.

Context facilitates performance on a classic cross-cultural emotion perception task.

The majority of studies designed to assess cross-cultural emotion perception use a choice-from-array task in which participants are presented with brief emotion stories and asked to choose between target and foil cues. This task has been widely criticized, evoking a lively and prolonged debate about whether it inadvertently helps participants to perform better than they otherwise would, resulting in the appearance of universality. In 3 studies, we provide a strong test of the hypothesis that the classic choice-from-array task constitutes a potent source of context that shapes performance. Participants from a remote small-scale (the Hadza hunter-gatherers of Tanzania) and 2 urban industrialized (China and the United States) cultural samples selected target vocalizations that were contrived for 6 non-English, nonuniversal emotion categories at levels significantly above chance. In studies of anger, disgust, fear, happiness, sadness, and surprise, above chance performance is interpreted as evidence of universality. These studies support the hypothesis that choice-from-array tasks encourage evidence for cross-cultural emotion perception. We discuss these findings with reference to the history of cross-cultural emotion perception studies, and suggest several processes that may, together, give rise to the appearance of universal emotions. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Cross-Species Evidence of Interplay Between Neural Connectivity at the Micro- and Macroscale of Connectome Organization in Human, Mouse, and Rat Brain.

The mammalian brain describes a multiscale system. At the microscale, axonal, dendritic, and synaptic elements ensure neuron-to-neuron communication, and at the macroscale, large-scale projections form the anatomical wiring for communication between cortical areas. Although it is clear that both levels of neural organization play a crucial role in brain functioning, their interaction is not extensively studied. Connectome studies of the mammalian brain in cat, macaque, and human have recently shown that regions with larger and more complex pyramidal cells to have more macroscale corticocortical connections. In this study, we aimed to further validate these cross-scale findings in the human, mouse, and rat brain. We combined neuron reconstructions from the NeuroMorpho.org neuroarchitecture database with macroscale connectivity data derived from connectome mapping by means of tract-tracing (rat, mouse) and in vivo diffusion magnetic resonance imaging (human). Across these three mammalian species, we show cortical variation in neural organization to be associated with features of macroscale connectivity, with cortical variation in neuronal complexity explaining significant proportions of cortical variation in the number of white matter projections of cortical areas. Our findings converge on the notion of a relationship between features of micro- and macroscale neural connectivity to form a central aspect of mammalian neural architecture.

Line-Drawn Scenes Provide Sufficient Information for Discrimination of Threat and Mere Negativity.

Previous work using color photographic scenes has shown that human observers are keenly sensitive to different types of threatening and negative stimuli and reliably classify them by the presence, and spatial and temporal directions of threat. To test whether such distinctions can be extracted from impoverished visual information, we used 500 line drawings made by hand-tracing the original set of photographic scenes. Sixty participants rated the scenes on spatial and temporal dimensions of threat. Based on these ratings, trend analysis revealed five scene categories that were comparable to those identified for the matching color photographic scenes. Another 61 participants were randomly assigned to rate the valence or arousal evoked by the line drawings. The line drawings perceived to be the most negative were also perceived to be the most arousing, replicating the finding for color photographic scenes. We demonstrate here that humans are very sensitive to the spatial and temporal directions of threat even when they must extract this information from simple line drawings, and rate the line drawings very similarly to matched color photographs. The set of 500 hand-traced line-drawing scenes has been made freely available to the research community: http://www.kveragalab.org/threat.html.

Salience Network Connectivity Modulates Skin Conductance Responses in Predicting Arousal Experience.

Individual differences in arousal experience have been linked to differences in resting-state salience network connectivity strength. In this study, we investigated how adding task-related skin conductance responses (SCR), a measure of sympathetic autonomic nervous system activity, can predict additional variance in arousal experience. Thirty-nine young adults rated their subjective experience of arousal to emotionally evocative images while SCRs were measured. They also underwent a separate resting-state fMRI scan. Greater SCR reactivity (an increased number of task-related SCRs) to emotional images and stronger intrinsic salience network connectivity independently predicted more intense experiences of arousal. Salience network connectivity further moderated the effect of SCR reactivity: In individuals with weak salience network connectivity, SCR reactivity more significantly predicted arousal experience, whereas in those with strong salience network connectivity, SCR reactivity played little role in predicting arousal experience. This interaction illustrates the degeneracy in neural mechanisms driving individual differences in arousal experience and highlights the intricate interplay between connectivity in central visceromotor neural circuitry and peripherally expressed autonomic responses in shaping arousal experience.

Multimodal analysis of cortical chemoarchitecture and macroscale fMRI resting-state functional connectivity.

The cerebral cortex is well known to display a large variation in excitatory and inhibitory chemoarchitecture, but the effect of this variation on global scale functional neural communication and synchronization patterns remains less well understood. Here, we provide evidence of the chemoarchitecture of cortical regions to be associated with large-scale region-to-region resting-state functional connectivity. We assessed the excitatory versus inhibitory chemoarchitecture of cortical areas as an ExIn ratio between receptor density mappings of excitatory (AMPA, M1 ) and inhibitory (GABAA , M2 ) receptors, computed on the basis of data collated from pioneering studies of autoradiography mappings as present in literature of the human (2 datasets) and macaque (1 dataset) cortex. Cortical variation in ExIn ratio significantly correlated with total level of functional connectivity as derived from resting-state functional connectivity recordings of cortical areas across all three datasets (human I: P = 0.0004; human II: P = 0.0008; macaque: P = 0.0007), suggesting cortical areas with an overall more excitatory character to show higher levels of intrinsic functional connectivity during resting-state. Our findings are indicative of the microscale chemoarchitecture of cortical regions to be related to resting-state fMRI connectivity patterns at the global system's level of connectome organization. Hum Brain Mapp 37:3103-3113, 2016. © 2016 Wiley Periodicals, Inc.

Threat perception after the Boston Marathon bombings: The effects of personal relevance and conceptual framing.

We examined how the Boston Marathon bombings affected threat perception in the Boston community. In a threat perception task, participants attempted to "shoot" armed targets and avoid shooting unarmed targets. Participants viewing images of the bombings accompanied by affectively negative music and text (e.g., "Terror Strikes Boston") made more false alarms (i.e., more errors "shooting" unarmed targets) compared to participants viewing the same images accompanied by affectively positive music and text (e.g., "Boston Strong") and participants who did not view bombing images. This difference appears to be driven by decreased sensitivity (i.e., decreased ability to distinguish guns from non-guns) as opposed to a more liberal bias (i.e., favouring the "shoot" response). Additionally, the more strongly affected the participant was by the bombings, the more their sensitivity was reduced in the negatively framed condition, suggesting that this framing was particularly detrimental to the most vulnerable individuals in the affected community.

Bridging Cytoarchitectonics and Connectomics in Human Cerebral Cortex.

The rich variation in cytoarchitectonics of the human cortex is well known to play an important role in the differentiation of cortical information processing, with functional multimodal areas noted to display more branched, more spinous, and an overall more complex cytoarchitecture. In parallel, connectome studies have suggested that also the macroscale wiring profile of brain areas may have an important contribution in shaping neural processes; for example, multimodal areas have been noted to display an elaborate macroscale connectivity profile. However, how these two scales of brain connectivity are related-and perhaps interact-remains poorly understood. In this communication, we combined data from the detailed mappings of early twentieth century cytoarchitectonic pioneers Von Economo and Koskinas (1925) on the microscale cellular structure of the human cortex with data on macroscale connectome wiring as derived from high-resolution diffusion imaging data from the Human Connectome Project. In a cross-scale examination, we show evidence of a significant association between cytoarchitectonic features of human cortical organization-in particular the size of layer 3 neurons-and whole-brain corticocortical connectivity. Our findings suggest that aspects of microscale cytoarchitectonics and macroscale connectomics are related. SIGNIFICANCE STATEMENT: One of the most widely known and perhaps most fundamental properties of the human cortex is its rich variation in cytoarchitectonics. At the same time, neuroimaging studies have also revealed cortical areas to vary in their level of macroscale connectivity. Here, we provide evidence that aspects of local cytoarchitecture are associated with aspects of global macroscale connectivity, providing insight into the question of how the scales of micro-organization and macro-organization of the human cortex are related.

Comparison of diffusion tractography and tract-tracing measures of connectivity strength in rhesus macaque connectome.

With the mapping of macroscale connectomes by means of in vivo diffusion-weighted MR Imaging (DWI) rapidly gaining in popularity, one of the necessary steps is the examination of metrics of connectivity strength derived from these reconstructions. In the field of human macroconnectomics the number of reconstructed fiber streamlines (NOS) is more and more used as a metric of cortico-cortical interareal connectivity strength, but the link between DWI NOS and in vivo animal tract-tracing measurements of anatomical connectivity strength remains poorly understood. In this technical report, we communicate on a comparison between DWI derived metrics and tract-tracing metrics of projection strength. Tract-tracing information on projection strength of interareal pathways was extracted from two commonly used macaque connectome datasets, including (1) the CoCoMac database of collated tract-tracing experiments of the macaque brain and (2) the high-resolution tract-tracing dataset of Markov and Kennedy and coworkers. NOS and density of reconstructed fiber pathways derived from DWI data acquired across 10 rhesus macaques was found to positively correlate to tract-tracing based measurements of connectivity strength across both the CoCoMac and Markov dataset (both P < 0.001), suggesting DWI NOS to form a valid method of assessment of the projection strength of white matter pathways. Our findings provide confidence of in vivo DWI connectome reconstructions to represent fairly realistic estimates of the wiring strength of white matter projections. Our cross-modal comparison supports the notion of in vivo DWI to be a valid methodology for robust description and interpretation of brain wiring.

Conceptualizing and experiencing compassion.

Does compassion feel pleasant or unpleasant? Westerners tend to categorize compassion as a pleasant or positive emotion, but laboratory compassion inductions, which present another's suffering, may elicit unpleasant feelings. Across two studies, we examined whether prototypical conceptualizations of compassion (as pleasant) differ from experiences of compassion (as unpleasant). After laboratory-based neutral or compassion inductions, participants made abstract judgments about compassion relative to various emotion-related adjectives, thereby providing a prototypical conceptualization of compassion. Participants also rated their own affective states, thereby indicating experiences of compassion. Conceptualizations of compassion were pleasant across neutral and compassion inductions. After exposure to others' suffering, however, participants felt increased levels of compassion and unpleasant affect, but not pleasant affect. After neutral inductions, participants reported more pleasant than unpleasant affect, with moderate levels of compassion. Thus, prototypical conceptualizations of compassion are pleasant, but experiences of compassion can feel pleasant or unpleasant. The implications for emotion theory in general are discussed.

Condoned or condemned: the situational affordance of anger and shame in the United States and Japan.

Two studies tested the idea that the situations that people encounter frequently and the situations that they associate most strongly with an emotion differ across cultures in ways that can be understood from what a culture condones or condemns. In a questionnaire study, N = 163 students from the United States and Japan perceived situations as more frequent to the extent that they elicited condoned emotions (anger in the United States, shame in Japan), and they perceived situations as less frequent to the extent that they elicited condemned emotions (shame in the United States, anger in Japan). In a second study, N = 160 students from the United States and Japan free-sorted the same situations. For each emotion, the situations could be organized along two cross-culturally common dimensions. Those situations that touched upon central cultural concerns were perceived to elicit stronger emotions. The largest cultural differences were found for shame; smaller, yet meaningful, differences were found for anger.

Dissociable large-scale networks anchored in the right anterior insula subserve affective experience and attention.

Meta-analytic summaries of neuroimaging studies point to at least two major functional-anatomic subdivisions within the anterior insula that contribute to the detection and processing of salient information: a dorsal region that is routinely active during attention tasks and a ventral region that is routinely active during affective experience. In two independent samples of cognitively normal human adults, we used intrinsic functional connectivity magnetic resonance imaging to demonstrate that the right dorsal and right ventral anterior insula are nodes in separable large-scale functional networks. Furthermore, stronger intrinsic connectivity within the right dorsal anterior insula network was associated with better performance on a task involving attention and processing speed whereas stronger connectivity within the right ventral anterior insula network was associated with more intense affective experience. These results support the hypothesis that the identification and manipulation of salient information is subserved by at least two brain networks anchored in the right anterior insula that exhibit distinct large-scale topography and dissociable behavioral correlates.