Dissociable modulation of overt visual attention in valence and arousal revealed by topology of scan path

Emotional stimuli have evolutionary significance for the survival of organisms; therefore, they are attention-grabbing and are processed preferentially. The neural underpinnings of two principle emotional dimensions in affective space, valence (degree of pleasantness) and arousal (intensity of evoked emotion), have been shown to be dissociable in the olfactory, gustatory and memory systems. However, the separable roles of valence and arousal in scene perception are poorly understood. In this study, we asked how these two emotional dimensions modulate overt visual attention. Twenty-two healthy volunteers freely viewed images from the International Affective Picture System (IAPS) that were graded for affective levels of valence and arousal (high, medium, and low). Subjects' heads were immobilized and eye movements were recorded by camera to track overt shifts of visual attention. Algebraic graph-based approaches were introduced to model scan paths as weighted undirected path graphs, generating global topology metrics that characterize the algebraic connectivity of scan paths. Our data suggest that human subjects show different scanning patterns to stimuli with different affective ratings. Valence salient stimuli (with neutral arousal) elicited faster and larger shifts of attention, while arousal salient stimuli (with neutral valence) elicited local scanning, dense attention allocation and deep processing. Furthermore, our model revealed that the modulatory effect of valence was linearly related to the valence level, whereas the relation between the modulatory effect and the level of arousal was nonlinear. Hence, visual attention seems to be modulated by mechanisms that are separate for valence and arousal.     

Meet OLAF,a Good Friend of the IAPS! The Open Library of Affective Foods: A Tool to Investigate the Emotional Impact of Food in Adolescents

In the last decades, food pictures have been repeatedly employed to investigate the emotional impact of food on healthy participants as well as individuals who suffer from eating disorders and obesity. However, despite their widespread use, food pictures are typically selected according to each researcher''s personal criteria, which make it difficult to reliably select food images and to compare results across different studies and laboratories. Therefore, to study affective reactions to food, it becomes pivotal to identify the emotional impact of specific food images based on wider samples of individuals. In the present paper we introduce the Open Library of Affective Foods (OLAF), which is a set of original food pictures created to reliably select food pictures based on the emotions they prompt, as indicated by affective ratings of valence, arousal, and dominance and by an additional food craving scale. OLAF images were designed to allow simultaneous use with affective images from the International Affective Picture System (IAPS), which is a well-known instrument to investigate emotional reactions in the laboratory. The ultimate goal of the OLAF is to contribute to understanding how food is emotionally processed in healthy individuals and in patients who suffer from eating and weight-related disorders. The present normative data, which was based on a large sample of an adolescent population, indicate that when viewing affective non-food IAPS images, valence, arousal, and dominance ratings were in line with expected patterns based on previous emotion research. Moreover, when viewing food pictures, affective and food craving ratings were consistent with research on food cue processing. As a whole, the data supported the methodological and theoretical reliability of the OLAF ratings, therefore providing researchers with a standardized tool to reliably investigate the emotional and motivational significance of food. The OLAF database is publicly available at zenodo.org.     

Cognitive Alexithymia Is Associated with the Degree of Risk for Psychosis

Alexithymia is a personality construct denoting emotion processing problems. It has been suggested to encompass two dimensions: a cognitive and affective dimension. The cognitive dimension is characterized by difficulties in identifying, verbalizing and analyzing emotions, while the affective dimension reflects the level of emotional arousal and imagination. Alexithymia has been previously proposed as a risk factor for developing psychosis. More specifically, the two alexithymia dimensions might be differentially related to the vulnerability for psychosis. Therefore, we examined the two dimensions of alexithymia, measured with the BVAQ in 94 siblings of patients with schizophrenia, 52 subjects at ultra-high risk (UHR) for developing psychosis, 38 patients with schizophrenia and 109 healthy controls. The results revealed that siblings and patients had higher levels of cognitive alexithymia compared to controls. In addition, subjects at UHR for psychosis had even higher levels of cognitive alexithymia compared to the siblings. The levels of affective alexithymia in siblings and patients were equal to controls. However, UHR individuals had significantly lower levels of affective alexithymia (i.e. higher levels of emotional arousal and fantasizing) compared to controls. Alexithymia was further related to subclinical levels of negative and depressive symptoms. These findings indicate that alexithymia varies parametrically with the degree of risk for psychosis. More specifically, a type-II alexithymia pattern, with high levels of cognitive alexithymia and normal or low levels of affective alexithymia, might be a vulnerability factor for psychosis.     

Affective responses by adults with autism are reduced to social images but elevated to images related to circumscribed interests

Individuals with autism spectrum disorders (ASD) demonstrate increased visual attention and elevated brain reward circuitry responses to images related to circumscribed interests (CI), suggesting that a heightened affective response to CI may underlie their disproportionate salience and reward value in ASD. To determine if individuals with ASD differ from typically developing (TD) adults in their subjective emotional experience of CI object images, non-CI object images and social images, 213 TD adults and 56 adults with ASD provided arousal ratings (sensation of being energized varying along a dimension from calm to excited) and valence ratings (emotionality varying along dimension of approach to withdrawal) for a series of 114 images derived from previous research on CI. The groups did not differ on arousal ratings for any image type, but ASD adults provided higher valence ratings than TD adults for CI-related images, and lower valence ratings for social images. Even after co-varying the effects of sex, the ASD group, but not the TD group, gave higher valence ratings to CI images than social images. These findings provide additional evidence that ASD is characterized by a preference for certain categories of non-social objects and a reduced preference for social stimuli, and support the dissemination of this image set for examining aspects of the circumscribed interest phenotype in ASD.     

The Influence of a Working Memory Task on Affective Perception of Facial Expressions

In a dual-task paradigm, participants performed a spatial location working memory task and a forced two-choice perceptual decision task (neutral vs. fearful) with gradually morphed emotional faces (neutral ∼ fearful). Task-irrelevant word distractors (negative, neutral, and control) were experimentally manipulated during spatial working memory encoding. We hypothesized that, if affective perception is influenced by concurrent cognitive load using a working memory task, task-irrelevant emotional distractors would bias subsequent perceptual decision-making on ambiguous facial expression. We found that when either neutral or negative emotional words were presented as task-irrelevant working-memory distractors, participants more frequently reported fearful face perception - but only at the higher emotional intensity levels of morphed faces. Also, the affective perception bias due to negative emotional distractors correlated with a decrease in working memory performance. Taken together, our findings suggest that concurrent working memory load by task-irrelevant distractors has an impact on affective perception of facial expressions.     

Mapping the semantic space for the subjective experience of emotional responses to odors

Two studies were conducted to examine the nature of the verbal labels that describe emotional effects elicited by odors. In Study 1, a list of terms selected for their relevance to describe affective feelings induced by odors was assessed while participants were exposed to a set of odorant samples. The data were submitted to a series of exploratory factor analyses to 1) reduce the set of variables to a smaller set of summary scales and 2) get a preliminary sense of the differentiation of affective feelings elicited by odors. The goal of Study 2 was to replicate the findings of Study 1 with a larger sample of odorant samples and participants and to validate the preliminary model obtained in Study 1 by using confirmatory factor analysis. Overall, the findings point to a structure of affective responses to odors that differs from the classical taxonomies of emotion such as posited by discrete or bidimensional emotion theories. These findings suggest that the subjective affective experiences or feelings induced by odors are structured around a small group of dimensions that reflect the role of olfaction in well-being, social interaction, danger prevention, arousal or relaxation sensations, and conscious recollection of emotional memories.     

Sensitivity to reward loss as an indicator of animal emotion and welfare

The scientific study of animal emotion is an important emerging discipline in subjects ranging from neuroscience to animal welfare research. In the absence of direct measures of conscious emotion, indirect behavioural and physiological measures are used. However, these may have significant limitations (e.g. indicating emotional arousal but not valence (positivity versus negativity)). A new approach, taking its impetus from human studies, proposes that biases in information processing, and underlying mechanisms relating to the evaluation of reward gains and losses, may reliably reflect emotional valence in animals. In general, people are more sensitive to reward losses than gains, but people in a negative affective state (e.g. depression) are particularly sensitive to losses. This may underlie broader findings such as an enhanced attention to, and memory of, negative events in depressed individuals. Here we show that rats in unenriched housing, who typically exhibit indicators of poorer welfare and a more negative affective state than those in enriched housing, display a prolonged response to a decrease in anticipated food reward, indicating enhanced sensitivity to reward loss. Sensitivity to reward reduction may thus be a valuable new indicator of animal emotion and welfare.     

Dynamic emotional and neural responses to music depend on performance expression and listener experience

Apart from its natural relevance to cognition, music provides a window into the intimate relationships between production, perception, experience, and emotion. Here, emotional responses and neural activity were observed as they evolved together with stimulus parameters over several minutes. Participants listened to a skilled music performance that included the natural fluctuations in timing and sound intensity that musicians use to evoke emotional responses. A mechanical performance of the same piece served as a control. Before and after fMRI scanning, participants reported real-time emotional responses on a 2-dimensional rating scale (arousal and valence) as they listened to each performance. During fMRI scanning, participants listened without reporting emotional responses. Limbic and paralimbic brain areas responded to the expressive dynamics of human music performance, and both emotion and reward related activations during music listening were dependent upon musical training. Moreover, dynamic changes in timing predicted ratings of emotional arousal, as well as real-time changes in neural activity. BOLD signal changes correlated with expressive timing fluctuations in cortical and subcortical motor areas consistent with pulse perception, and in a network consistent with the human mirror neuron system. These findings show that expressive music performance evokes emotion and reward related neural activations, and that music's affective impact on the brains of listeners is altered by musical training. Our observations are consistent with the idea that music performance evokes an emotional response through a form of empathy that is based, at least in part, on the perception of movement and on violations of pulse-based temporal expectancies.     

Self-Reported Trait Mindfulness and Affective Reactivity: A Motivational Approach Using Multiple Psychophysiological Measures

As a form of attention, mindfulness is qualitatively receptive and non-reactive, and is thought to facilitate adaptive emotional responding. One suggested mechanism is that mindfulness facilitates disengagement from an affective stimulus and thereby decreases affective reactivity. However, mindfulness has been conceptualized as a state, intervention, and trait. Because evidence is mixed as to whether self-reported trait mindfulness decreases affective reactivity, we used a multi-method approach to study the relationship between individual differences in self-reported trait mindfulness and electrocortical, electrodermal, electromyographic, and self-reported responses to emotional pictures. Specifically, while participants (N = 51) passively viewed pleasant, neutral, and unpleasant IAPS pictures, we recorded high-density (128 channels) electrocortical, electrodermal, and electromyographic data to the pictures as well as to acoustic startle probes presented during the pictures. Afterwards, participants rated their subjective valence and arousal while viewing the pictures again. If trait mindfulness spontaneously reduces general emotional reactivity, then for individuals reporting high rather than low mindfulness, response differences between emotional and neutral pictures would show relatively decreased early posterior negativity (EPN) and late positive potential (LPP) amplitudes, decreased skin conductance responses, and decreased subjective ratings for valence and arousal. High mindfulness would also be associated with decreased emotional modulation of startle eyeblink and P3 amplitudes. Although results showed clear effects of emotion on the dependent measures, in general, mindfulness did not moderate these effects. For most measures, effect sizes were small with rather narrow confidence intervals. These data do not support the hypothesis that individual differences in self-reported trait mindfulness are related to spontaneous emotional responses during picture viewing.     

The Cognitive Building Blocks of Emotion Regulation: Ability to Update Working Memory Moderates the Efficacy of Rumination and Reappraisal on Emotion

The ability to regulate emotions is a critical component of healthy emotional functioning. Therefore, it is important to determine factors that contribute to the efficacy of emotion regulation. The present article examined whether the ability to update emotional information in working memory is a predictor of the efficacy of rumination and reappraisal on affective experience both at the trait level (Study 1) and in daily life (Study 2). In both studies, results revealed that the relationship between use of reappraisal and high arousal negative emotions was moderated by updating ability. Specifically, use of reappraisal was associated with decreased high arousal negative emotions for participants with high updating ability, while no significant relationship was found for those with low updating ability. In addition, both studies also revealed that the relationship between rumination and high arousal negative emotions was moderated by updating ability. In general, use of rumination was associated with elevated high arousal negative emotions. However, this relationship was blunted for participants with high updating ability. That is, use of rumination was associated with less elevated high arousal negative emotions for participants with high updating ability. These results identify the ability to update emotional information in working memory as a crucial process modulating the efficacy of emotion regulation efforts.     

Interplay between Affect and Arousal in Recognition Memory

Background

Emotional states linked to arousal and mood are known to affect the efficiency of cognitive performance. However, the extent to which memory processes may be affected by arousal, mood or their interaction is poorly understood.

Methodology/Principal Findings

Following a study phase of abstract shapes, we altered the emotional state of participants by means of exposure to music that varied in both mood and arousal dimensions, leading to four different emotional states: (i) positive mood-high arousal; (ii) positive mood-low arousal; (iii) negative mood-high arousal; (iv) negative mood-low arousal. Following the emotional induction, participants performed a memory recognition test. Critically, there was an interaction between mood and arousal on recognition performance. Memory was enhanced in the positive mood-high arousal and in the negative mood-low arousal states, relative to the other emotional conditions.

Conclusions/Significance

Neither mood nor arousal alone but their interaction appears most critical to understanding the emotional enhancement of memory.     

Human implicit memory for irrelevant dimension values is similar to rats' incidental memory in simultaneous discrimination tasks

Participants completed a category-learning task in which they needed to discover which of three stimulus dimensions (shape, color or size) was relevant. After meeting a learning criterion (nine of 10 consecutive correct responses), participants continued making categorization choices and response latencies associated with these trials were examined. In both Experiments 1 and 2, people responded reliably faster when correct responses matched the previous responses with respect to irrelevant dimension values. Thus, they demonstrated a form of incidental short-term memory analogous to that we previously reported in studies of rats. In Experiment 2, participants' explicit memory for irrelevant dimension values was assessed after category learning was complete. The results indicated that people were unaware of the irrelevant dimension values encountered on trials preceding surprise probe trials. This indicates that memory for the irrelevant dimension values was implicit (i.e. unconscious). The findings are discussed with respect to both human and non-human studies of hippocampus-independent memory and implicit memory.     

A Diffusion Model Analysis of Decision Biases Affecting Delayed Recognition of Emotional Stimuli

Previous empirical work suggests that emotion can influence accuracy and cognitive biases underlying recognition memory, depending on the experimental conditions. The current study examines the effects of arousal and valence on delayed recognition memory using the diffusion model, which allows the separation of two decision biases thought to underlie memory: response bias and memory bias. Memory bias has not been given much attention in the literature but can provide insight into the retrieval dynamics of emotion modulated memory. Participants viewed emotional pictorial stimuli; half were given a recognition test 1-day later and the other half 7-days later. Analyses revealed that emotional valence generally evokes liberal responding, whereas high arousal evokes liberal responding only at a short retention interval. The memory bias analyses indicated that participants experienced greater familiarity with high-arousal compared to low-arousal items and this pattern became more pronounced as study-test lag increased; positive items evoke greater familiarity compared to negative and this pattern remained stable across retention interval. The findings provide insight into the separate contributions of valence and arousal to the cognitive mechanisms underlying delayed emotion modulated memory.     

Conditions under which Arousal Does and Does Not Elevate Height Estimates

We present a series of experiments that explore the boundary conditions for how emotional arousal influences height estimates. Four experiments are presented, which investigated the influence of context, situation-relevance, intensity, and attribution of arousal on height estimates. In Experiment 1, we manipulated the environmental context to signal either danger (viewing a height from above) or safety (viewing a height from below). High arousal only increased height estimates made from above. In Experiment 2, two arousal inductions were used that contained either 1) height-relevant arousing images or 2) height-irrelevant arousing images. Regardless of theme, arousal increased height estimates compared to a neutral group. In Experiment 3, arousal intensity was manipulated by inserting an intermediate or long delay between the induction and height estimates. A brief, but not a long, delay from the arousal induction served to increase height estimates. In Experiment 4, an attribution manipulation was included, and those participants who were made aware of the source of their arousal reduced their height estimates compared to participants who received no attribution instructions. Thus, arousal that is attributed to its true source is discounted from feelings elicited by the height, thereby reducing height estimates. Overall, we suggest that misattributed, embodied arousal is used as a cue when estimating heights from above that can lead to overestimation.     

Affective modulation of multiple memory systems

The hippocampus and caudate nucleus are anatomical components of relatively independent memory systems and recent research has focused on the nature of the interaction between these two systems. The amygdala exerts a general modulatory influence on memory storage processes related, in part, to an organism's level of affective or emotional arousal. Moreover, affective state can influence the use of different memory systems, and the amygdala may mediate this effect of emotion on memory. Recent evidence indicates that the amygdala modulates the separate types of memory mediated by the hippocampus and caudate nucleus. Recent human brain imaging studies also point to both sex- and hemisphere-related asymmetries in amygdala participation in emotionally influenced memory.     

Effects of unconscious processing on implicit memory for fearful faces

Emotional stimuli can be processed even when participants perceive them without conscious awareness, but the extent to which unconsciously processed emotional stimuli influence implicit memory after short and long delays is not fully understood. We addressed this issue by measuring a subliminal affective priming effect in Experiment 1 and a long-term priming effect in Experiment 2. In Experiment 1, a flashed fearful or neutral face masked by a scrambled face was presented three times, then a target face (either fearful or neutral) was presented and participants were asked to make a fearful/neutral judgment. We found that, relative to a neutral prime face (neutral-fear face), a fearful prime face speeded up participants' reaction to a fearful target (fear-fear face), when they were not aware of the masked prime face. But this response pattern did not apply to the neutral target. In Experiment 2, participants were first presented with a masked faces six times during encoding. Three minutes later, they were asked to make a fearful/neutral judgment for the same face with congruent expression, the same face with incongruent expression or a new face. Participants showed a significant priming effect for the fearful faces but not for the neutral faces, regardless of their awareness of the masked faces during encoding. These results provided evidence that unconsciously processed stimuli could enhance emotional memory after both short and long delays. It indicates that emotion can enhance memory processing whether the stimuli are encoded consciously or unconsciously.     

Valence and Arousal Ratings for 420 Finnish Nouns by Age and Gender

Language-and culture-specific norms are needed for research on emotion-laden stimuli. We present valence and arousal ratings for 420 Finnish nouns for a sample of 996 Finnish speakers. Ratings are provided both for the whole sample and for subgroups divided by age and gender in light of previous research suggesting age- and gender-specific reactivity to the emotional content in stimuli. Moreover, corpus-based frequency values and word length are provided as objective psycholinguistic measures of the nouns. The relationship between valence and arousal mainly showed the curvilinear relationship reported in previous studies. Age and gender effects on valence and arousal ratings were statistically significant but weak. The inherent affective properties of the words in terms of mean valence and arousal ratings explained more of the variance in the ratings. In all, the findings suggest that language- and culture-related factors influence the way affective properties of words are rated to a greater degree than demographic factors. This database will provide researchers with normative data for Finnish emotion-laden and emotionally neutral words. The normative database is available in Database S1.     

Evidence that disgust evolved to protect from risk of disease

Disgust is a powerful human emotion that has been little studied until recently. Current theories do not coherently explain the purpose of disgust, nor why a wide range of stimuli can provoke a similar emotional response. Over 40 000 individuals completed a web-based survey using photo stimuli. Images of objects holding a potential disease threat were reported as significantly more disgusting than similar images with little or no disease relevance. This pattern of response was found across all regions of the world. Females reported higher disgust sensitivity than males; there was a constant decline in disgust sensitivity over the life course; and the bodily fluids of strangers were found more disgusting than those of close relatives. These data provide evidence that the human disgust emotion may be an evolved response to objects in the environment that represent threats of infectious disease.     

Analysis of elicited EEG synchronization and desynchronization in emotional activation: temporal and topographic characteristics

Event-related synchronization (ERS) and desynchronization (ERD) in delta, theta1, theta2, alpha1, alpha2, beta1, beta2, beta3, and gamma were measured in 20 healthy right-handed subjects in response to IAPS stimuli with low, moderate, and high arousal reactions. The 62-channel EEG was simultaneously recorded while subjects viewed sequentially presented pictures and subjectively rated them after each presentation. The results show that emotionally loaded stimuli induced higher ERS in the delta, theta1, theta2, beta1, beta3, and gamma bands along with combined ERD and ERS effects in alpha2 band. As to hemispheric asymmetries, the effects of emotional arousal were restricted not only to right parietal (theta1 and theta2 ERS, alpha2 ERD) but also to left frontal (theta2 ERS) regions. In terms of affective chronometry, lower theta was the first to catch the affective salience of incoming stimuli (time window 0-600 ms after the stimulus input). For theta2, alpha2, and gamma bands this process was delayed to 600-1000 ms.     

Effects of aversive stimuli on prospective memory. An event-related fMRI study

Prospective memory (PM) describes the ability to execute a previously planned action at the appropriate point in time. Although behavioral studies clearly showed that prospective memory performance is affected by the emotional significance attributed to the intended action, no study so far investigated the brain mechanisms subserving the modulatory effect of emotional salience on PM performance. The general aim of the present study was to explore brain regions involved in prospective memory processes when PM cues are associated with emotional stimuli. In particular, based on the hypothesised critical role of the prefrontal cortex in prospective memory in the presence of emotionally salient stimuli, we expected a stronger involvement of aPFC when the retrieval and execution of the intended action is cued by an aversive stimulus. To this aim BOLD responses of PM trials cued by aversive facial expressions were compared to PM trials cued by neutral facial expressions. Whole brain analysis showed that PM task cued by aversive stimuli is differentially associated with activity in the right lateral prefrontal area (BA 10) and in the left caudate nucleus. Moreover a temporal shift between the response of the caudate nucleus that preceded that of aPFC was observed. These findings suggest that the caudate nucleus might provide an early analysis of the affective properties of the stimuli, whereas the anterior lateral prefrontal cortex (BA10) would be involved in a slower and more deliberative analysis to guide goal-directed behaviour.