The regulation of breathing assumes the presence of certain long-enduring properties of the atmosphere and the respiratory system. Vision assumes the presence of certain evolutionarily stable properties of surfaces, objects, and terrestrial spectral distributions. The lactase digestive enzyme presupposes an infant diet of milk with lactose. And each emotion presupposes that certain cues signal the presence of a structure of events and conditions that held true during the evolution of that emotion. Disgust circuits presume a world in which rotten smells signal toxins or microbial contamination, for example.
Accordingly, to understand an adaptation as a problem-solver, one needs to model the enduring properties of the task-environment that selected for that adaptation -- the environment of evolutionary adaptedness, or EEA. Although the hominid line is thought to have first differentiated from the chimpanzee lineage on the African savannas and woodlands, the EEA is not a place or time. Thus, statistical regularities define the EEA for any given adaptation. The conditions that characterize the EEA are usefully decomposed into a constellation of specific environmental regularities that had a systematic though not necessarily unvarying impact on reproduction and that endured long enough to work evolutionary change on the design of an adaptation.
These regularities can include complex conditionals e. Conceptualizing the EEA in statistical terms is fundamental to the functional definition of emotion that we presented above and will elucidate below. The cognitive science resolution of the mind-body problem. Evolutionary psychology starts with a fundamental insight from cognitive psychology: The brain is a machine designed to process information. From this perspective, one can define the mind as a set of information-processing procedures cognitive programs that are physically embodied in the neural circuitry of the brain.
For cognitive scientists, brain and mind are terms that refer to the same system, which can be described in two complementary ways - either in terms of its physical properties the neural , or in terms of its information-processing operation the mental. This approach allows mental operations to be described with great precision: one is led to specify what information is extracted from the environment, what procedures act to transform it, what formats are involved in its representation or storage, and what operations access it to govern decision-making, physiological or behavioral regulation, or further information integration.
Also, because it provides an intelligible way of relating physical and mental phenomena, discoveries in brain science e. An evolutionary perspective makes clear why the cognitive or computational level of description is more than an analogy. Whereas other parts of the body were designed for lifting loads, grinding food, chemically extracting nutrients, and so on, the brain was designed by evolution to use information derived from the environment and the body to functionally regulate behavior and the body. The brain came into existence and, over evolutionary time, accreted its present complex structure because, in ancestral populations, mutations that created or altered cognitive programs such that they more successfully carried out adaptively consequential information-processing tasks were differentially retained, replicated, and incorporated into our species' neural design.
The ancestral world posed recurrent information-processing problems, such as What substances are best to eat? Information-processing programs - food preferences and aversions, or rules for inferring emotions from facial expressions - acquired one set of design features rather than many others because the retained features better computed solutions to these information-processing problems.
Over evolutionary time, it was the computational properties of alternative neural circuits - their relative ability to solve adaptive information-processing tasks - that caused some neural circuits to be selected for, and others to be selected out. A key task for psychologists, then, is to discover, inventory and map the "circuit logic" of the collection of programs that constitute the human mind, and relate how that adaptive logic maps onto the suite of informational problems faced by our hunter-gatherer ancestors.
Emotion and computation. It may strike some as odd to speak about love or jealousy or disgust in computational terms. In everyday language, the term "cognition" is often used to refer to a particular subset of information-processing - roughly, the effortful, conscious, voluntary, deliberative kind of thinking one does when solving a mathematics problem or playing chess: what is sometimes called "cold cognition". This use of "cognition" falls out of the folk psychological classification of thinking as distinct from feeling and emotion, and it appears in a few subfields of psychology as well particularly those concerned with education and the acquisition of skills that must be explicitly taught.
As a result, one sometimes sees articles in the psychological literature on how emotion, affect, or mood influence "cognition". However, from an evolutionary cognitive perspective, one cannot sensibly talk about emotion affecting cognition because cognition refers to a language for describing all of the brain's operations, including emotions and reasoning whether deliberative or nonconscious , and not to any particular subset of operations. If the brain evolved as a system of information-processing relations, then emotions are, in an evolutionary sense, best understood as information-processing relations - i.
Initially, the commitment to exploring the underlying computational architecture of the emotions may strike one as odd or infelicitous, but it leads to a large number of scientific payoffs, as we will sketch out below. Thus, the claim that emotion is computational does not mean that an evolutionary psychological approach reduces the human experience to bloodless, affectless, disembodied ratiocination. Every mechanism in the brain - whether it does something categorizable as "cold cognition" such as inducing a rule of grammar or judging a probability or as "hot cognition" such as computing the intensity of parental fear, the imperative to strike an adversary, or an escalation in infatuation - depends on an underlying computational organization to give its operation its patterned structure, as well as a set of neural circuits to physically implement it.
Of course, shifting terminology e. But an evolutionary and computational view of emotion can open up for exploration new empirical possibilities obscured by other frameworks. An evolutionary perspective breaks categories like "thinking" into a large set of independent domain-specific programs, and so opens up the possibility that distinct emotions affect separate inference programs in diverse, yet functionally patterned, ways, rather than in a single, aggregate way.
Domain specificity and functional specialization. A basic engineering principle is that the same machine is rarely capable of solving two different problems equally well. Cork-screws and cups have different properties because they are solutions to different problems, and each therefore solves its targeted problem better than the other. Similarly, natural selection has constructed different tissues and organs, like the heart for pumping blood and the liver for detoxifying poisons, for exactly this reason. This same principle applies to our evolved cognitive programs and neural circuitry.
Different information-processing problems usually require different procedures for their successful solution. For example, to solve the adaptive problem of selecting a good mate, one's choices must be guided by qualitatively different standards than when choosing the right food, or the right habitat, or the right meaning for an unfamiliar word.
Emotion and Adaptation by Richard S. Lazarus
Speed, reliability, and efficiency can be engineered into specialized mechanisms because they do not need to make trade-offs between mutually incompatible task demands, and because they can use problem-solving principles that work in one domain but not in others. The application of these principles to the design of the mind has convinced many scientists, including most evolutionary psychologists, that the human cognitive architecture is multimodular : that it is composed of a large number of information-processing programs, many of which are functionally specialized for solving a different adaptive problem.
These adaptations appear to be domain-specific expert systems, equipped with "crib sheets": inference procedures, regulatory rules, motivational priorities, goal-definitions, and assumptions that embody knowledge, regulatory structure, and value weightings specific to an evolved problem domain. These generate correct or, at least, adaptive outputs that would not be warranted on the basis of perceptual data processed through some general-purpose decisional algorithm.
These domain-specific inference systems have a distinct advantage over domain-independent ones, akin to the difference between experts and novices: experts can solve problems faster and more efficiently than novices because they already know a lot about the problem domain, and because they are equipped with specialized tools and practices. Each adaptive problem recurred millions of times in the EEA, and so manifested a statistical and causal structure whose elements were available for specialized exploitation by design features of the evolving adaptation.
For example, predators use darkness and cover to ambush. Physical appearance varies with fertility and health. Children one's mother regularly feeds are usually genetic siblings. Such specializations, by embodying "innate knowledge" about the problem space, operate better than any general learning strategy could. A child did not have to wait to be ambushed and killed in the dark to prudently modulate her activities. Adults did not need to observe the negative effects of incest, because the Westermarck mechanism mobilizes disgust towards having sex with likely siblings Shepher Selection detects the individually unobservable.
Animals subsist on information. The single most limiting resource to reproduction is not food or safety or access to mates, but what makes them each possible: the information required for making adaptive behavioral choices. However, many important features of the world cannot be perceived directly. Cognitive adaptations can use perceivable events as cues for inferring the status of important, nonperceivable sets of conditions, provided there was a predictable probabilistic relationship between them that was maintained over evolutionary time.
It does this by testing randomly generated alternative designs, each of which embodies different assumptions about the structure of the world, and retaining the ones that succeed most effectively. The most effective design will be the one that best embodies design features that reflect most closely the actual long-term statistical structure of the ancestral world. Designs whose features exploited these real but ontogenetically unobservable relationships outperformed those that depended on different relationships, or that only responded to conditions the individual could observe during her lifetime.
For example, the negative effects of incestuous conceptions are difficult for any individual to observe in the absence of a modern controlled study with numerous participants, much less integrate rationally into one's motivational system. Fortunately, the consequences of incest over evolutionary time selected for specialized disgust mechanisms that reflected the ancestral distribution of choice-consequence pairings, and so are designed to guide humans away from incestuous unions between fertile adults, given appropriate cues of familial connection such as co-residence in the first years of life Shepher, Evolved psychological adaptations are selected to use cues that 1 can be reliably and easily detected by the individual, and 2 reliably predicted the hidden structure of conditions relevant to determining which course of action one should take.
The functional structure of an emotion program evolved to match the evolutionarily summed structure of its target situation. The set of human emotion programs assumed their evolved designs through interacting with the statistically defined structure of human environments of evolutionary adaptedness. Each emotion program was constructed by a selective regime imposed by a particular evolutionarily recurrent situation.
By an evolutionarily recurrent situation, we mean a cluster of repeated probabilistic relationships among events, conditions, actions, and choice-consequences, that endured over a sufficient stretch of evolutionary time to have had selective consequences on the design of the mind, and that were probabilistically associated with cues detectable by humans. For example, the condition of having a mate plus the condition of one's mate copulating with someone else constitutes a situation of sexual infidelity: a situation that has recurred over evolutionary time, even though it has not happened to every individual.
Associated with this situation were cues reliable enough to allow the evolution of a "situation detector" e. Even more importantly, there were many necessarily or probabilistically associated elements that tended to be present in the situation of infidelity as encountered among our hunter-gatherer ancestors.
Additional elements include: 1 a sexual rival with a capacity for social action and violence, as well as allies of the rival; 2 a discrete probability that one's mate has conceived with the sexual rival; 3 changes in the net lifetime reproductive returns of investing further in the mating relationship; 4 a probable decrease in the degree to which the unfaithful mate's mechanisms value the victim of infidelity the presence of an alternative mate lowers replacement costs ; 5 a cue that the victim of the infidelity will likely have been deceived about a range of past events, leading the victim to confront the likelihood that his or her memory is permeated with false information; 6 the victim's status and reputation for being effective at defending his or her interests in general would be likely to plummet, inviting challenges in other arenas.
These are just a few of the many factors that constitute a list of elements associated in a probabilistic cluster, and that constitute the evolutionary recurrent structure of a situation of sexual infidelity. The emotion of sexual jealousy evolved in response to these properties of the world, and there should be evidence of this in its computational design.
Emotion programs have evolved to take such elements into account, whether they can be perceived or not. Thus, not only do cues of a situation trigger an emotion mode, but embedded in that emotion mode is a way of seeing the world and feeling about the world related to the ancestral cluster of associated elements. Depending on the intensity of the jealousy evoked, less and less evidence will be required for an individual to believe that these conditions apply to their personal situation.
Individuals with morbid jealousy, for example, may hallucinate counterfactual but evolutionarily thematic contents. To the extent that situations exhibit a structure repeated over evolutionary time, their statistical properties will be used as the basis for natural selection to build an emotion program whose detailed design features are tailored for that situation.
This is accomplished by selection, acting over evolutionary time, differentially incorporating program components that dovetail with individual items on the list of properties probabilistically associated with the situation. For example, if in ancestral situations of sexual infidelity there was a substantially higher probability of a violent encounter than in its absence, then the sexual jealousy program will have been shaped by the distillation of those encounters, and the jealousy subroutines will have been adjusted to prepare for violence in proportion to the raised probability in the ancestral world.
Natural selection acts too slowly to have updated the mind to post-hunter-gatherer conditions. Each of these sub-elements and the adaptive circuits they require can be added together to form a general theory of sexual jealousy. It is the relationship between the summed details of the ancestral condition and the detailed structure of the resulting emotion program that makes this approach so useful for emotion researchers.
Each functionally distinct emotion state - fear of predators, guilt, sexual jealousy, rage, grief, and so on - will correspond to an integrated mode of operation that functions as a solution designed to take advantage of the particular structure of the recurrent situation or triggering condition to which that emotion corresponds. This approach can be used to create theories of each individual emotion, through three steps: 1 Reconstructing the clusters of properties of ancestral situations; 2 Constructing engineering analyses about how each of the known or suspected psychological mechanisms in the human mental architecture should be designed to deal with each ancestral condition or cluster of conditions, and integrating these into a model of the emotion program; 3 Constructing or conducting experiments and other investigations to test, and revise the models of emotion programs.
It is also important to understand that evolutionarily recurrent situations can be arrayed along a spectrum in terms of how rich or skeletal is the set of probabilistically associated elements that defines the situation. A richly structured situation, such as sexual infidelity or predator ambush will support a richly substructured emotion program in response to the many ancestrally correlated features: Many detailed adjustments will be made to many psychological mechanisms as instructions for the mode of operation.
In contrast, some recurrent situations have less structure i. For example, surges of happiness or joy are an emotion program that evolved to respond to the recurrent situation of encountering unexpected positive events as will be explained. The class of events captured by "unexpectedly positive" is extremely broad, general, and have only a few additional properties in common.
Emotion programs at the most general and skeletal end of this spectrum correspond to what some call "mood" happiness, sadness, excitement, anxiety, playfulness, homesickness, and so on. To characterize an emotion adaptation, one must identify the following properties of environments and of mechanisms. Examples of situations are being in a depleted nutritional state, competing for maternal attention, being chased by a predator, being about to ambush an enemy, having few friends, experiencing the death of a spouse, being sick, having experienced a public success, having others act in a way that damages you without regard for your welfare, having injured a valued other through insufficient consideration or self-other behavioral trade-offs; having a baby.
For example, what is the best course of action when others take the products of your labor without your consent? What is the best course of action when you are in a depleted nutritional state? What is the best course of action when a sibling makes a sexual approach? A multimodular mind must be full of "demons": algorithms that detect situations. The New Hacker's Dictionary defines a "demon" as a "portion of a program that is not invoked explicitly, but that lies dormant waiting for some condition s to occur. Situation-detecting subprograms lie dormant, until they are activated by a specific constellation of cues that precipitates the analysis of whether a particular ancestral situation has arisen.
If the assessment is positive, it sends the signal that activates the associated emotion program. Emotion demons need two kinds of subroutines:. They take the cues in 3 as input. Animals should be designed to "detect" what situation they are in on the basis of cues, stored variables, and specialized interpretation algorithms. Selection will not shape decision rules so that they act solely on the basis of what is most likely to be true, but rather on the basis of the weighted consequences of acts given that something is held to be true.
Should you walk under a tree that might conceal a predator? The benefits of calories saved via a shortcut, scaled by the probability that there is no predator in the tree, must be weighed against the benefits of avoiding becoming catfood, scaled by the probability that there is a predator in the tree. Because the costs and benefits of false alarms, misses, hits, and correct rejections are often unequal, the decision rules may still treat as true situations that are unlikely to be true.
In the modern world, this behavior may look "irrational" as is the case with many phobias , but we do it because such decisions were adaptive under ancestral conditions. Situation detecting algorithms can be of any degree of complexity, from demons that monitor single cues e. Inherent in this approach is the expectation that the human mind has a series of evolved subsystems designed to represent events in terms of evolutionarily recurrent situations and situational subcomponents.
The operation of these representational systems are not necessarily consciously accessible. By their structure, they impose an evolutionary organization on representational spaces that are updated by data inputs. When the representational space assumes certain configurations, an interpretation is triggered that activates the associated emotion program - corresponding approximately to what others have called a cognitive appraisal see, e. It is important to recognize that the evolutionary past frames the experienced present, because these situation-detecting algorithms provide the dimensions and core elements out of which many cross-culturally recurring representations of the world are built.
To some extent, the world we inhabit is shaped by the continuous interpretive background commentary provided by these mechanisms. The prioritizing algorithms define which emotion modes are compatible e. Depending on the relative importance of the situations and the reliability of the cues, the prioritizing algorithms decide which emotion modes to activate and deactivate, and to what degree.
Selection, through ancestral mutant experiments, would have sorted emotions based on the average importance of the consequences stemming from each, and the extent to which joint activation was mutually incompatible or facilitating. Prioritizing algorithms can be thought of as a supervisory system operating over all of the emotions.
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In addition, information is fed back into the emotion program from other programs and systems that assess body states, which may govern the intensity, trajectory, supplantation, or termination of the emotion. Some modes of activation of the cognitive system are accompanied by a characteristic feeling state, a certain quality of experience. The fact that we are capable of becoming aware of certain physiological states - our hearts thumping, bowels evacuating, stomach tightening - is surely responsible for some of the qualia evoked by emotion states that entrain such responses.
The fact that we are capable of becoming aware of certain mental states - such as retrieved memories of past events - is probably responsible for other qualia. But it is also possible that in some cases, the characteristic feeling state that accompanies an emotion mode results in part from mechanisms that allow us to sense the signal that activates and deactivates the relevant programs.
Such internal sensory mechanisms - a kind of cognitive proprioception - can be selected for if there are mechanisms that require as input the information that a particular emotion mode has been activated. This might be true, for example, of mechanisms designed to inhibit certain stimulus driven actions when the conditions are not auspicious. For example, there should be algorithms in the auditory system that, upon detecting the fear signal see 6 , reset signal detection thresholds, increasing acuity for predator-relevant sounds.
Any controllable biological process that, by shifting its performance in a specifiable way, would lead to enhanced average fitness outcomes should have come to be partially governed by emotional state see 7 above. Such processes include:. Goals : The cognitive mechanisms that define goal-states and choose among goals in a planning process should be influenced by emotions.
For example, vindictiveness-a specialized subcategory of anger-may define "injuring the offending party" as a goal state to be achieved. Although the evolved functional logic of this process is deterrence, this function need not be represented, either consciously or unconsciously, by the mechanisms that generate the vindictive behavior. Motivational priorities : Mechanisms involved in hierarchically ranking goals or calibrating other kinds of motivational and reward systems should be emotion-dependent.
What may be extremely unpleasant in one state, such as harming another, may seem satisfying in another state e. Different evolutionarily recurrent situations predict the presence - visible or invisible - of different opportunities, risks, and payoffs, so motivational thresholds and valences should be entrained. For example, a loss of face should increase the motivation to take advantage of opportunities for status-advancement, and decrease attention to attendant costs.
Information-gathering motivations : Because establishing which situation you are in has enormous consequences for the appropriateness of behavior, the process of detection should in fact involve specialized inference procedures and specialized motivations to discover whether certain suspected facts are true or false. What one is curious about, what one finds interesting, what one is obsessed with discovering should all be emotion-specific. Imposed conceptual frameworks : Emotions should prompt construals of the world in terms of concepts that are appropriate to the decisions that must be made.
When angry, domain-specific concepts such as social agency, fault, responsibility and punishment will be assigned to elements in the situation. When hungry, the food-nonfood distinction will seem salient. When endangered, safety-categorization frames will appear. The world will be carved up into categories based partly on what emotional state an individual is in.
Perceptual mechanisms : Perceptual systems may enter emotion-specific modes of operation. When fearful, acuity of hearing may increase. Specialized perceptual inference systems may be mobilized as well: If you've heard rustling in the bushes at night, human and predator figure-detection may be particularly boosted, and not simply visual acuity in general.
In fact, non-threat interpretations may be depressed, and the same set of shadows will "look threatening"-that is, given a specific threatening interpretation such as "a man with a knife"-or not, depending on emotion-state. Memory : The ability to call up particularly appropriate kinds of information out of long term memory ought to be influenced.
A woman who has just found strong evidence that her husband has been unfaithful may find herself flooded by a torrent of memories about small details that seemed meaningless at the time but that now fit into an interpretation of covert activity. We also expect that what is stored about present experience will also be differentially regulated.
Important or shocking events, for example, may be stored in great detail as has been claimed about "flashbulb memories", for example , but other more moderate emotion-specific effects may occur as well. Attention : The entire structure of attention, from perceptual systems to the contents of high level reasoning processes, should be regulated by emotional state. Positive emotions may broaden attentional focus Fredrickson, Physiology : Each organ system, tissue, or process is a potential candidate for emotion-specific regulation, and "arousal" is insufficiently specific to capture the detailed coordination involved.
Each emotion program should send out a different pattern of instructions to the face and limb muscles, the autonomic system, etc. Changes in circulatory, respiratory, and gastrointestinal functioning are well-known and documented, as are changes in endocrinological function. We expect thresholds regulating the contraction of various muscle groups to change with certain emotion states, reflecting the probability that they will need to be employed. Similarly, immune allocation and targeting may vary with disgust, with the potential for injury, or with the demands of extreme physical exertion.
Communication and emotional expressions : Emotion programs are expected to mobilize many emotion-specific effects on the subcomponents of the human psychological architecture relevant to communication. Most notably, many emotion programs produce characteristic species-typical displays that broadcast to others the emotion state of the individual Ekman, Ekman and his colleagues have established in a careful series of landmark studies that many emotional expressions are human universals, both generated and recognized reliably by humans everywhere they have been tested Ekman, Indeed, many emotional expressions appear to be designed to be informative, and these have been so reliably informative that humans co-evolved automated interpreters of facial displays of emotion that decode these public displays into knowledge of others' mental states.
It is surely true that people sometimes "lie" with their faces. But programs for inferring emotion states from facial displays would not have evolved unless doing so created a net advantage for the inferer, suggesting that these inferences were warranted more often than not. Two things are communicated by an authentic emotional expression: 1 that the associated emotion program has been activated in an individual, providing observers with information about the state of that individual's mental programs and physiology e.
Both are highly informative, and emotional expressions provide a continuous commentary on the underlying meaning of things to companions. This provokes the question: Why did selection build facial, vocal and postural expressions at all? More puzzlingly, why are they often experienced as automatic and involuntary? From an evolutionary perspective, sometimes it is beneficial to provide information to others, and other times it is injurious, so most evolved communication systems involve close regulation of whether to transmit information or not. Usually this leads to a system, like language, in which the decision to communicate something or not can be made by the individual in detailed response to the immediate circumstances.
The apparent selective disadvantages of honestly and automatically broadcasting one's emotional state have led Fridlund , for example, to argue that expressions must be voluntary and intentional communications largely unconnected to emotion state. Undoubtedly they sometimes are. But even when people deliberately lie, microexpressions of face and voice often leak out Ekman, , suggesting that certain emotion programs do in fact create involuntarily emitted signals that reliably broadcast the person's emotion state. Natural selection has shaped emotion programs to signal their activation, or not, on an emotion by emotion basis.
Summing for each emotion program considered by itself jealousy, loneliness, disgust, predatoriness, parental love, sexual attraction, gratitude, fear , there was a net benefit or cost to having others know that mental state, averaged across individuals over evolutionary time.
For those recurrent situations in which, on average, it was beneficial to share one's emotion state and hence assessment of the situation with those one was with, species-typical facial and other expressions of emotion were constructed by selection. For example, fear was plausibly beneficial to signal, because it signaled the presence of a danger that might also menace one's kin and cooperators, and also informed others in a way that might recruit assistance.
Nevertheless, averaged over evolutionary time, it was functional for the organism to signal the activation of only some emotion states. The conditions favoring signaling an emotion are hard to meet, so only some emotions out of the total species-typical set are associated with distinctive, species-typical facial expressions. There should be a larger set of emotions that have no automatic display. Jealousy, guilt, or boredom are all genuine emotions lacking distinctive signals.
This changes the question from: Why are emotions automatically signaled? When they are denied or repressed, or when they overflow without control, they have a negative effect on our personal and relational health. At the human level, each emotion represents the satisfaction of a need and it constitutes a way of adaptive emotional intelligence, with its own and independent circuits from the neuronal circuits of cognitive intelligence, as shown by Daniel Goleman. Between these two emotions is located the fight or flight response, the most primitive of our adaptive strategies geared towards survival.
The emotion of sadness produces the internalization of the feeling and the possibility of the encounter with oneself. Obsessions and compulsions related to the need of security and confidence regarding our thoughts and actions represent the embryo of discipline, vocation and consecration which favors the self fulfillment. Furthermore, actors were never presented in precisely overlapping retinotopic locations comparable to e.
Together, these results rule out other possible low-level or post-perceptual explanations for emotional action aftereffects that might account for some other demonstrations of high-level perceptual aftereffects. We cannot, however, rule out the contribution of adaptation in motion processing mechanisms, which is likely to occur simultaneously when viewing complex action stimuli e.
Action speed is an important cue in the emotion recognition from body actions e. Other postural and dynamic features that do not rely on speed are also critical for expression and for perception of emotional actions e. These postural e. It is therefore plausible that the aftereffects observed here resulted from adaptation at multiple levels of action visual processing. Importantly, our studies with emotional action aftereffects indicate that identity plays a role in the representations of emotional actions, and identity can modulate the way that emotional actions are processed.
When the identities of the adapting and test actors were the same aftereffects were larger than when the identities of the adapting and test actors were different. Furthermore, when the identities of the adapting and test actors were the same, the aftereffects did not appear to decline over the time period we tested up to In contrast, when the identities of the adapting and test actors were different, the aftereffects declined over time. Thus, emotional action aftereffects show different magnitudes and decay functions dependent upon the relationship between the identity of adapting and test actors.
One explanation for our results is that emotional actions are represented in both an identity-dependent and identity-independent way. When adapting and test actors have the same identity, the aftereffects increase in magnitude and appear to last much longer than typical high-level action or face aftereffects.
The increase in aftereffect magnitude could be explained by an increase in visual similarity between the adapting and test stimuli. However, this increase in visual similarity could not abolish the decay of the aftereffect over time in the way that we observe here. A parsimonious explanation is that an identity-dependent mechanism is additionally adapted when adapting and test stimuli have the same identities, resulting in a larger aftereffect and a longer-lasting influence on the representations of the emotions of individual actors.
Such long-lasting aftereffects are not necessarily unusual, as they have been previously reported with other social stimuli such as gaze direction e. These previous effects, however, were observed with much longer adaptation periods than we have used in our experiment reported here. Identity has also been shown to affect the decay of face aftereffects Kiani et al. For both identity-dependent and identity-independent aftereffects, adaptation never transferred across different actions.
Different actions have different kinematics, and bodily expression of emotion is dependent upon the characteristic kinematics of the action e. The failure of adaptation to one set of characteristic movements to exert an influence on the perception of a very different set of movements, suggests that actor emotion may be coded within action specific neural mechanisms. In some ways, these results parallel findings of face emotion aftereffects. Evidence from face adaptation studies suggest the existence of a common visual representation that underlies the coding of face identity and face expression Ellamil et al.
This common representation may contain one dimension that codes both the identity and expression, and the other dimension that is selective for identity or expression Calder, ; Rhodes et al. These findings are also consistent with Haxby et al. Specifically, the fusiform face area FFA , predominantly involved in the coding of the invariant aspects of the face, may also play a supportive role in the coding of emotional expressions Haxby et al. Similarly, processing of voice identity and voice emotion is thought to be processed by seperate, but interacting systems Belin et al. While early analysis of vocal input from primary auditory cortex to the middle part of the STS is shared for different types of vocal information, at the highest level of voice processing the pathways become independent Belin et al.
The similarity between the aftereffects we observed with emotional action adaptation and our understanding of how faces Haxby et al. Grossman et al. The neural substrates underlying our ability to recognize actor identity from body form and motion have not been well delineated, but may involve the Extrastriate Body Area EBA; Saxe et al.
Adaptation in more anterior regions of the temporal cortex that takes into account actor identity may underlie the emotional action aftereffect that is influenced by the identity of the individual actors; although this suggestion is more speculative.
Emotion and Adaptation
Anterior cortical regions do not show the same adaptation characteristics as the more posterior cortical regions cf. Verhoef et al. The less characteristic longer lasting adaptation effects observed here, and also seen in some face adaptation experiments e. Such long lasting aftereffects may shift the observer's reference point according to their recent experience in order to optimize the sensory processing of the external world over a longer period of time.
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This study has demonstrated that prior exposure to emotional whole-body actions influences the perception of the emotion conveyed by subsequent actions via a perceptual adaptation mechanism. Our visual adaptation experiments reveal two separate processing mechanisms for emotional actions with different characteristics: one mechanism that processes actor emotion irrespective of actor identity, and one that processes actor emotion taking into account actor identity.
This organization parallels recent data on the processing of face information that suggest that rather than completely separate processing of emotion and identity, representations of emotion, and identity can interact. These mechanisms we study here would not only help us to determine the emotions of individuals around us from their actions and behavior, but also critically ensure the identity of the individual is linked to the specific emotions expressed.
We thank Andy Young and Bruce Keefe for comments on an earlier draft of this manuscript. All data supporting this study are provided as supplementary information accompanying this paper. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
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