Theoretical,and epistemological challenges in scientific investigations of complex emotional states in animals

This review brings to light critical epistemological and theoretical considerations when studying complex emotional states in animals. We discuss anthropomorphic and Umwelt perspectives of nonhuman animals and the ways in which distinct theories of consciousness and neural processing may restrict the potential for the development of knowledge on the topic. Within the same line of argumentation, we consider influences of the debate between monism and dualism and psychology’s behaviorism and cognitive theories. Finally, we contrast the affective consciousness, higher-order emotional consciousness, and constructed emotion theories to further our understanding of complex emotional states in animals.     

When can we choose to forget? An ERP study into item-method directed forgetting of emotional words

Emotionally arousing information is treated in a specialised manner across a number of different processing stages, and memory for affective events is often found to be heightened by virtue of this. However, in some cases, emotional experiences might be the very ones that we would like to forget. Here, two item-method directed forgetting studies are presented which investigate people's ability to intentionally forget affective words when stimuli and memory instructions are presented simultaneously. In the first experiment an interaction between task instruction and emotional content was evident in a diminished directed forgetting effect for emotional words, suggesting that they may be relatively resistant to deliberate forgetting. The interaction between instruction and emotion appeared both in free recall of words and in a yes-no recognition task. In the second study, an ERP procedure was utilised to investigate whether emotion modulates the effects of instruction during the initial encoding of stimuli. Recognition data again showed a clear interaction between instruction and emotion, with a reduced directed forgetting effect for emotional words. The ERP data demonstrated evidence for individual effects of both emotion and instruction during encoding; however, despite this, no evidence for an interaction between these factors was evident in the ERP data. As such, we conclude that even when study items are presented simultaneously with their associated memory instructions, neither does emotion prevent differential processing of directed forgetting instruction, nor does memory instruction prevent differential processing of emotion during early encoding. Implications are discussed in relation to the directed forgetting literature and more broadly with respect to circumstances under which emotion and cognitive processing work in parallel or in competition with each other.     

情绪自动化加工的证据与争议

情绪信息对人类的生存非常重要, 研究者一直在探讨情绪产生的本质问题：情绪的产生是“自动”的吗？已有的大量实验证据表明, 情绪可以被快速识别, 甚至在非注意条件下或者无意识条件下, 情绪信息也可以被大脑(尤其是杏仁核)识别。大量实验结果分析了情绪自动化加工的表现, 并探讨了情绪自动化加工的大脑机制：皮层下通路。视觉信息在视网膜接受后, 经上丘脑与枕核传递至杏仁核, 这条通路被称为皮层下通路, 负责情绪信息的快速化加工。但是新的研究对已有的实验范式与技术手段进行了质疑, 反驳情绪的自动化加工, 并质疑皮层下通路存在的可能性。为了进一步理清情绪加工的本质与大脑机制, 今后的研究应当采用更为严格地控制意识、注意资源的实验范式, 以及采用更精确、更高时间与空间分辨率的技术手段。     

Emotion and Pain: A Functional Cerebral Systems Integration

Emotion and pain are psychological constructs that have received extensive attention in neuropsychological research. However, neuropsychological models of emotional processing have made more progress in describing how brain regions interact to process emotion. Theories of emotional processing can describe inter-hemispheric and intra-hemispheric interactions during emotional processing. Due to similarities between emotion and pain, it is thought that emotional models can be applied to pain. The following review examines the neuropsychology of emotion and pain using a functional cerebral systems approach. Specific comparisons are made between pain and anger. Attention is given to differences in cerebral function and physiology that may contribute to the processing of emotion and pain. Suggestions for future research in emotion and pain are given.     

Gender differences in implicit and explicit processing of emotional facial expressions as revealed by event-related theta synchronization

Emotion information processing may occur in 2 modes that are differently represented in conscious awareness. Fast online processing involves coarse-grained analysis of salient features and is not represented in conscious awareness; offline processing takes hundreds of milliseconds to generate fine-grained analysis and is represented in conscious awareness. These processing modes may be studied using event-related electroencephalogram theta synchronization as a marker of emotion processing. Two experiments were conducted that differed on the mode of emotional information presentation. In the explicit mode, subjects were explicitly instructed to evaluate the emotional content of presented stimuli; in the implicit mode, their attention was directed to other features of the stimulus. In the implicit mode, theta synchronization was most pronounced in the early processing stage, whereas in the explicit mode, it was more pronounced in the late processing stage. The early processing stage was more pronounced in men, whereas the late processing stage was more pronounced in women. Implications of these gender differences in emotion processing for well-documented differences in social behavior are discussed.     

道德判断中的情绪因素——从认知神经科学的角度进行探讨

道德判断中的情绪与认知因素是社会认知神经科学研究的重要议题。本文对近年来社会认知神经科学领域中有关情绪影响道德判断的研究发现和相关理论进行了回顾和总结。介绍了外源性情绪影响道德判断的行为研究证据,大脑情绪功能障碍者在道德判断中的异常表现以及道德两难任务所对应的大脑激活模式,着重探讨了情绪因素影响道德判断的神经机制和道德判断中情绪加工与认知加工相互作用的过程。揭示了情绪是道德判断的必要因素,道德判断是情绪加工和认知加工协同作用的结果,并在此基础上提出了该领域未来发展的设想。     

Modulation of retrieval processing reflects accuracy of emotional source memory

There is considerable evidence that encoding and consolidation of memory are modulated by emotion, but the retrieval of emotional memories is not well characterized. Here we manipulated the emotional context with which affectively neutral stimuli were associated during encoding, allowing us to examine neural activity associated with retrieval of emotional memories without confounding the emotional attributes of cue items and the retrieved context. Using a source memory procedure we were also able to compare how retrieval processing was modulated when the emotional encoding context was recollected or not. An interaction between emotional encoding context and accuracy of source memory revealed that successful retrieval of emotional context was associated with activity in left amygdala, and a left frontotemporal network including anterior insula, prefrontal cortex and cingulate. In contrast, when contextual retrieval was unsuccessful, items encoded in emotional contexts elicited enhanced activity in right amygdala and a right-lateralized network that included extrastriate visual areas. These findings indicate distinct effects of emotion on successful and unsuccessful retrieval of source information, including lateralization of amygdala responses.     

Mere social categorization modulates identification of facial expressions of emotion

The ability of the human face to communicate emotional states via facial expressions is well known, and past research has established the importance and universality of emotional facial expressions. However, recent evidence has revealed that facial expressions of emotion are most accurately recognized when the perceiver and expresser are from the same cultural ingroup. The current research builds on this literature and extends this work. Specifically, we find that mere social categorization, using a minimal-group paradigm, can create an ingroup emotion-identification advantage even when the culture of the target and perceiver is held constant. Follow-up experiments show that this effect is supported by differential motivation to process ingroup versus outgroup faces and that this motivational disparity leads to more configural processing of ingroup faces than of outgroup faces. Overall, the results point to distinct processing modes for ingroup and outgroup faces, resulting in differential identification accuracy for facial expressions of emotion.     

Emotion and consciousness

Consciousness and emotion feature prominently in our personal lives, yet remain enigmatic. Recent advances prompt further distinctions that should provide more experimental traction: we argue that emotion consists of an emotion state (functional aspects, including emotional response) as well as feelings (the conscious experience of the emotion), and that consciousness consists of level (e.g. coma, vegetative state and wakefulness) and content (what it is we are conscious of). Not only is consciousness important to aspects of emotion but structures that are important for emotion, such as brainstem nuclei and midline cortices, overlap with structures that regulate the level of consciousness. The intersection of consciousness and emotion is ripe for experimental investigation, and we outline possible examples for future studies.     

Positive emotion modulates cognitive control: An event‐related potentials study

There is substantial evidence to indicate that negative emotion can modulate cognitive control processing. However, only a few studies have investigated this effect with positive emotion. Therefore, the present study explored the electrophysiological correlates of the impact of positive emotional stimuli on cognitive control, using event‐related potentials (ERPs). Sixteen healthy young adults completed a modified Simon task (Simon, 1969). Behavioral data indicated that reaction times were faster for positive emotional stimuli than for neutral emotional stimuli in the incongruent condition, but not in the congruent condition, which suggested that positive emotion expedited conflict resolution. The ERP data showed that two negative ERP components (N300–400 and N450–550) were associated with the positive emotional stimuli in the incongruent minus congruent condition. It is suggested that these components may respectively be related to the conflict monitoring (N300–400) and response selection (N450–550) stages of cognitive control processing. Overall, our results indicated that positive emotion could facilitate cognitive control processing. These results are in line with the neuropsychological theory, according to which, positive emotion could modulate cognitive control mediated by increased dopamine levels in frontal brain areas.     

Processing orientation and emotion recognition

There is evidence that some emotional expressions are characterized by diagnostic cues from individual face features. For example, an upturned mouth is indicative of happiness, whereas a furrowed brow is associated with anger. The current investigation explored whether motivating people to perceive stimuli in a local (i.e., feature-based) rather than global (i.e., holistic) processing orientation was advantageous for recognizing emotional facial expressions. Participants classified emotional faces while primed with local and global processing orientations, via a Navon letter task. Contrary to previous findings for identity recognition, the current findings are indicative of a modest advantage for face emotion recognition under conditions of local processing orientation. When primed with a local processing orientation, participants performed both significantly faster and more accurately on an emotion recognition task than when they were primed with a global processing orientation. The impacts of this finding for theories of emotion recognition and face processing are considered.     

Sad or fearful? The influence of body posture on adults’ and children’s perception of facial displays of emotion

The current research investigated the influence of body posture on adults' and children's perception of facial displays of emotion. In each of two experiments, participants categorized facial expressions that were presented on a body posture that was congruent (e.g., a sad face on a body posing sadness) or incongruent (e.g., a sad face on a body posing fear). Adults and 8-year-olds made more errors and had longer reaction times on incongruent trials than on congruent trials when judging sad versus fearful facial expressions, an effect that was larger in 8-year-olds. The congruency effect was reduced when faces and bodies were misaligned, providing some evidence for holistic processing. Neither adults nor 8-year-olds were affected by congruency when judging sad versus happy expressions. Evidence that congruency effects vary with age and with similarity of emotional expressions is consistent with dimensional theories and "emotional seed" models of emotion perception.     

The valence-specific laterality effect in free viewing conditions: The influence of sex, handedness, and response bias

The right hemisphere has often been viewed as having a dominant role in the processing of emotional information. Other evidence indicates that both hemispheres process emotional information but their involvement is valence specific, with the right hemisphere dealing with negative emotions and the left hemisphere preferentially processing positive emotions. This has been found under both restricted (Reuter-Lorenz & Davidson, 1981) and free viewing conditions (Jansari, Tranel, & Adophs, 2000). It remains unclear whether the valence-specific laterality effect is also sex specific or is influenced by the handedness of participants. To explore this issue we repeated Jansari et al.'s free-viewing laterality task with 78 participants. We found a valence-specific laterality effect in women but not men, with women discriminating negative emotional expressions more accurately when the face was presented on the left-hand side and discriminating positive emotions more accurately when those faces were presented on the right-hand side. These results indicate that under free viewing conditions women are more lateralised for the processing of facial emotion than are men. Handedness did not affect the lateralised processing of facial emotion. Finally, participants demonstrated a response bias on control trials, where facial emotion did not differ between the faces. Participants selected the left-hand side more frequently when they believed the expression was negative and the right-hand side more frequently when they believed the expression was positive. This response bias can cause a spurious valence-specific laterality effect which might have contributed to the conflicting findings within the literature.     

Effortful versus automatic emotional processing in schizophrenia: Insights from a face-vignette task

Adaptive emotional responding relies on dual automatic and effortful processing streams. Dual-stream models of schizophrenia (SCZ) posit a selective deficit in neural circuits that govern goal-directed, effortful processes versus reactive, automatic processes. This imbalance suggests that when patients are confronted with competing automatic and effortful emotional response cues, they will exhibit diminished effortful responding and intact, possibly elevated, automatic responding compared to controls. This prediction was evaluated using a modified version of the face-vignette task (FVT). Participants viewed emotional faces (automatic response cue) paired with vignettes (effortful response cue) that signalled a different emotion category and were instructed to discriminate the manifest emotion. Patients made less vignette and more face responses than controls. However, the relationship between group and FVT responding was moderated by IQ and reading comprehension ability. These results replicate and extend previous research and provide tentative support for abnormal conflict resolution between automatic and effortful emotional processing predicted by dual-stream models of SCZ.     

The experience of positive emotion is associated with the automatic processing of positive emotional words

The current study examines the relationship between attention bias for positive emotional words and self-reported emotional experience. Previous research suggests that the experience of positive emotion momentarily broadens cognitive processes, potentially allowing individuals to build an array of enduring personal resources. However, it is unknown whether the experience of positive emotion also broadens emotional information processing. Participants included 60 healthy undergraduate students who completed measures of psychopathology, self-reported emotional experience, and an emotional Stroop task designed to measure attentional bias to positive and negative emotional information. Results indicate significant associations between reaction times for high-intensity happiness words and self-reported high levels of positive emotion and low levels of negative emotions. These associations were not present for low intensity happiness words. Findings suggest that individuals who experience high levels of positive emotion and low levels of negative emotion demonstrate an attention bias for positive information and, from an information processing perspective, provide insight into the manner in which positive emotions broaden cognitive processes.     

Neural correlates of adaptive decision making for risky gains and losses

Do decisions about potential gains and potential losses require different neural structures for advantageous choices? In a lesion study, we used a new measure of adaptive decision making under risk to examine whether damage to neural structures subserving emotion affects an individual's ability to make adaptive decisions differentially for gains and losses. We found that individuals with lesions to the amygdala, an area responsible for processing emotional responses, displayed impaired decision making when considering potential gains, but not when considering potential losses. In contrast, patients with damage to the ventromedial prefrontal cortex, an area responsible for integrating cognitive and emotional information, showed deficits in both domains. We argue that this dissociation provides evidence that adaptive decision making for risks involving potential losses may be more difficult to disrupt than adaptive decision making for risks involving potential gains. This research further demonstrates the role of emotion in decision competence.     

情绪的自动加工与控制加工

由于人类注意资源有限,对相关刺激的加工经常以抑制对其它刺激的加工为代价。大量的研究表明,情绪的加工是注意加工过程的一个特例。通常认为情绪信息的加工是自动的,不需要注意资源的调节。但最近的研究结果倾向于支持对立的观点,即情绪信息的加工需要注意的控制。该文结合认知神经科学研究证据,对情绪的自动加工的观点提出质疑。主要的证据来自对情绪的皮层和皮层下通路的研究以及对盲视病人的研究。     

面孔情绪识别过程中的多通道效应

摘 要 面孔情绪识别过程中的多感觉通道效应指参与刺激加工的各个通道对面孔表情认知的综合影响。研究者们通过行为实验、事件相关电位以及脑成像技术等对该过程中的多个获取信息的通道进行研究,肢体表情、情绪性声音、特定气味能系统地影响面孔表情的情绪识别。一系列的研究对多通道效应的作用时间、潜在作用机制、相关激活脑区进行了探索。未来的研究可以整合脑网络的技术,并结合其他学科的新技术以更细致具体地考察这些通道下信息的物理属性所起的作用。 关键词 面孔表情 多通道效应 面孔情绪识别 肢体表情 情绪性声音 嗅觉信号     

Positive emotion impedes emotional but not cognitive conflict processing

Cognitive control enables successful goal-directed behavior by resolving a conflict between opposing action tendencies, while emotional control arises as a consequence of emotional conflict processing such as in irony. While negative emotion facilitates both cognitive and emotional conflict processing, it is unclear how emotional conflict processing is affected by positive emotion (e.g., humor). In 2 EEG experiments, we investigated the role of positive audiovisual target stimuli in cognitive and emotional conflict processing. Participants categorized either spoken vowels (cognitive task) or their emotional valence (emotional task) and ignored the visual stimulus dimension. Behaviorally, a positive target showed no influence on cognitive conflict processing, but impeded emotional conflict processing. In the emotional task, response time conflict costs were higher for positive than for neutral targets. In the EEG, we observed an interaction of emotion by congruence in the P200 and N200 ERP components in emotional but not in cognitive conflict processing. In the emotional conflict task, the P200 and N200 conflict effect was larger for emotional than neutral targets. Thus, our results show that emotion affects conflict processing differently as a function of conflict type and emotional valence. This suggests that there are conflict- and valence-specific mechanisms modulating executive control.     

Differentiating emotional processing and attention in psychopathy with functional neuroimaging

Individuals with psychopathy are often characterized by emotional processing deficits, and recent research has examined the specific contexts and cognitive mechanisms that underlie these abnormalities. Some evidence suggests that abnormal features of attention are fundamental to emotional deficits in persons with psychopathy, but few studies have demonstrated the neural underpinnings responsible for such effects. Here, we use functional neuroimaging to examine attention–emotion interactions among incarcerated individuals (n = 120) evaluated for psychopathic traits using the Hare Psychopathy Checklist–Revised (PCL-R). Using a task designed to manipulate attention to emotional features of visual stimuli, we demonstrate effects representing implicit emotional processing, explicit emotional processing, attention-facilitated emotional processing, and vigilance for emotional content. Results confirm the importance of considering mechanisms of attention when evaluating emotional processing differences related to psychopathic traits. The affective-interpersonal features of psychopathy (PCL-R Factor 1) were associated with relatively lower emotion-dependent augmentation of activity in visual processing areas during implicit emotional processing, while antisocial-lifestyle features (PCL-R Factor 2) were associated with elevated activity in the amygdala and related salience network regions. During explicit emotional processing, psychopathic traits were associated with upregulation in the medial prefrontal cortex, insula, and superior frontal regions. Isolating the impact of explicit attention to emotional content, only Factor 1 was related to upregulation of activity in the visual processing stream, which was accompanied by increased activity in the angular gyrus. These effects highlight some important mechanisms underlying abnormal features of attention and emotional processing that accompany psychopathic traits.