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The Nature of Emotions Human emotions have deep evolutionary roots, a fact that may explain their complexity and provide tools for clinical practice Robert Plutchik Imost everyone agrees that the study of emotion is one of the most confused (and sill open) chapters in the history of psychology. By one estimate, ‘more than 90 definitions of “emotion” were proposed over the course of the 20th century. If there is litle consensus (on the meaning of the erm, itis no wore der that there is much disagreement among contemporary theoreticians con- cerning the best way to conceptualize emotion and interprets role in life Tn everyday human existence we con- ceive of an emotion—anger, despair joy, srief—asa feeling, an inner state. The in ternal experience of emotion is highly personal and often confusing, paricular- ly because several emotions may be ex- perienced at the same time. Imagine, then, how dificult the objective study of ‘emotion must be. Mos of us often censor ‘our own thoughts and feelings, and we have leamed to be cautious about ac- cepting other people's comments about their feelings. The empirical study of a psychological phenomenon so complex and so elaborately cloaked cannot help butpresent.a special challenge. Compounding the distrust of verbal reports of emotion are the influences of behaviorism and psychoanalysis on psy- chological research. The behaviorists of the 20th century believed that the only truly reliable, objective information ob- tainable from living creatures was infor- mation about their behavior. A classical Robert Patch poisor emeritus atthe Albert Ensen Colle of Medicine and adj at the University of South Farida. He recived is PheD. from Columbia Univers. He he eure or contort more tha 260 ates, 5 chapters ‘and eight book ad heed sever bos. His, researc inet nclude testy of tions, ici and violence andthe sty of. spy proces. Ades for Pitch 4505 Der CreckBoulenard, Sarscta, FI. 34238, Internat: probanhome com 344 American Scientist Volume 8° ‘behaviorist would hold that emotion is an inner state and thus simply outside the realm of science, For their pact psy- choanalysts have made us aware that emotions may be repressed, inhibited or unconscious, and thus unavailable to in- trospection. Finally, language itself intro- duces ambiguity and does not make it easy to describe mixed emotions in an ‘unequivocal way. The meaning of emo- tion termsis often obscure. For example, ‘many people are not sure about the dif. ferences between fear and anxiety, guilt and shame, or envy and jealousy: As a result we often resort to metaphor to at- tempt to describe emotion, Think, for ex- ample, of such expressions as “blowing off steam,” “hating someone’s guts,” “pain inthe neck,” “lump in the throat” and “abroken heart.” How, then, can emotion be studied and understood? The challenge of devel- oping a theoretical approach is impor- tant, because emotions are an essential part of who we are and how we survive; ‘emotional distress impels people to seek help, and indeed the primary concern of psychotherapy is the repair of emotional disorders. To simply declare emotion ‘outside the bounds of scientific study ‘would be imresponsible. Ibelieve that a scientific andl therapeu tically useful understanding of emotions is possible In fact, there are several sci- entific intellectual traditions that have ealt with this issue. There are an evalue tionary (aunched by Charles Darwin), a psychophysiological (William James), a neurological (Walter Cannon) and a psy- chodynamic tradition (Sigmunal Freud), in addition to the cognitive perspective that began emerging in the 1950s. More recently neurcbiclogical evidence has be- gun to inform the discussion; however, identifying the structures ofthe brain re~ lated to emotion isnot a theory of emo- tion, nor can such a theory be built from, a knoviledge of the chemicals involved in mood states, just as an adequate theo- 1y of depression cannot be constructed simply from a knowledge of the avail ability ofserotonin. As the Univesity of Towa neuroscientist Antonio Damasio has pointed out, when the amassing of data does not resolve a complex issue it ray be necessary to find new ways to conceptualize the problem, 1 my view evolutionary theory pro- vides a way to unify a number of theo- retical perspectives. Using the tools and methods of evolutionary biology, and paling together information from other species, we can put emotions in a fune- tional ramewerk—define them in terms of what their adaptive function mightbe, and thus understand better theirbiclogi- cal basis and the apparent connections between them. Some work long theselines has been widely popularized in recent years: BY now we'veall heard authors of best-ell- ing books describe jealousy, love, am- ty and fear in dogs, cats, chimpanzees, baboons, elephants and lions. The bes” tiaries ofthe medieval period contained detailed descriptions of emotions in ani- als. The popular appeal of such expla- nations may lien their ability to touch a deep-seated sense ofthe connectedness ofall living things. Although many psy chologists have warmed of the dangers of anthropomorphism, recent thinking by cognitive scientists and others sees this atitude as an outdated prejudice The Rockefeller University zoologist Donald Griffin, one of the founders of the field of animal cognition, believes that the “charge of anthropomorphism is a conceited claim that only our species is capable of even the simplest conscious thinking,” But there is dan- ger in oversimplifying: A sophisticated understanding is eeded to inform clin- ical practice. Over the past four decades Thave pulled together evidence from various oe ‘Cosel Feer Aol re Figure Great egrets in breeding plumage fightin midair, displaying the agonistic behavior that is the precursor of human emotions suchas fea, anger and jealous. Human emotions, sys the author are best viewed though an evolutionary lens, as adaptations triggered by the challenges of studies to form a psychoevolutionary theory of emotion, withthe goals of cla- fying what emotions are, finding ways to measure them, relating emotions to ‘other psychological disciplines, and in- forming the practice of psychotherapy. ‘Like many concepts in science, emotions canbe best understood by making infer- ences from certain classes of evidence. Such inferences suggest that emotions or their evolutionary precursors (or proto- types) can be found among lower ani- snals as well as human beings, afact that ‘can provide fascinating evolutionary in- sights into our emotions, moods and personality traits. They suggest further ‘that emotion, cognition and action inter actin feedback loops andi that emotion canbe viewed ina structural model tied to adaptation, a ‘Survival and reprdlction that ae part of every organisms existence. An evolutionary approach, he argues, can sort ou the roles of emotion, impulse “and action and in a therapeutic seting, help people understand the cicumetances in which emotions can sometimes fil in their adaptive tasks, Evolution and Emotion What we call cognition—the activity of knowing, learning and thinking, of which emotion is a part—evolved over millions of years, Charles Darwin recog nized that the process of evolution by ‘natural selection applied not only to anatomic structures but also to an ani- mal’s “mind” and expressive behav {ora concsion that led him to writea book on emotional expression, Those ‘who have followed Darwin in studying the evolutionary origins of emotion have sought to understand how emo- fions increase evolutionary fitness for the individual ‘As mentioned above, a few evolu- tionary origins are easy to postulate. Fear ancl anxiety in people closely paral- lel the state of heightened arousal of an animal wo senses apredatorora threat twits offspring, a similarity thathas been found inreurechemica anatomical and ‘imaging studies that show these states are mediated by the limbic system, the part of the central nervous sytem com- ron to lower and higher animals, Love tnd emotional attachment cleazly pro mote pair bonding, reproduction and Jal investment basic to evoltion By fines in human beings, But the or- gins of some other emotions are harder to find. Is there a general principle that ante applied? “The place to start might be with the de finiion problem. An emotion isnot sim pif. feling ate. Emotion is a complex Eun of oesely connected evens that be fgns witha stinolus and includes feel- ings psychological changes, impulses to 2001 July-August 35 Figure 2. Microorganisms—here a parameciim, mixed algae diatoms and other plankton—take in food, excete waste products, avoid predators, reproduce, seek safe environments and explore their world. They must distinguish between predator and prey and, if their mode of reproduc tion requires i between a potential mate and a potential enemy. This information gathering and prediction and other universal adaptations found in all organisms constitute prototype ‘adaptations from which human emotion can be modeled. action and specific, goal-directed behav- for That is to say, feelings do nothappen {in isolation. They are responses to sig- nificant situations in an individual's life, and often they motivate actions. This de- nition of emotions allows the concept ‘tobe generalized to lower animals with- out difficulty. From his studies of ani mals, human infants and human adults, Darwin concluded that expressive be- Figure 3. Suckling her young, a suricate imeerkat mother keeps am eye on her sue roundings ina South African nature preserve, ready to protect her offspring from any ¢hreat. Peychologists believe cognitive functions evolved to serve emotional and biological ‘needs—in evolutionary tems, in order to pre- dict the future more effectively. In survival tuations cognitions and Inne states such as srousal tend tobe followed by impulses to 3 tion—perhaps, in this ease a defensive o pro teative posture attack or light 346 American Scents, Volume 89 haviors communicate information from ‘one animal to another about what is likely to happen; therefore they affect the chances of survival of the individual demonstrating the behavior. “Even in- sects,” he wrote in his 1872 book, “ex- [press anger, terror jealousy, and love by ‘heir stridulations.” Extending Darwin’s idea a bit, pro- pose that in general, emotions are acti- vated in an individual when issues of survival ae raised in factor by implica On. Such situations include threats, at- facks, poisonous substances or the sighting of a potential mate. The effect of the emotional state is to create an in- teraction between the individual and the event or stimulus that precipitated the emotion. The interaction usually takes the form of an attempt to reduce the disequilibrium and reestablish a state of comparative rest. Protozoologist Nicola Ricci of the University of Pisa in Ttaly pointed out {in 1990 that every single-celled organ- sm, from the blue-green alga to the eu- Karyote, isa complete, self-sufficient or- ganism, Single-celled organisms are exposed to daily risks in their environ ‘ments. They take in food, excrete waste products, avoid predators, reproduce by exchange of genes in many cases, seek safe environments and explore their microbiological world. Thus these simple organisms adapt to many of the same problems as higher, multicellular organisms. Bacteria are capable of very complex metabolic pathways, and, as, Ursula Goodenough detailed in these pages in 1991, viruses and bacteria have evolved tactics of camouflage, dis- traction and mimicry. Even plant cells such as green algae show defensive re- actions fo touch, and chemical mes- sages signal everything from alarm to sexual attraction in organisms from bacteria to human beings. ‘As one moves up the evolutionary ladder, it is remarkable to note that a small number of developmental genes can radically alter the behavior of cells and change an amoeba into a multi- celled organism. Developmental biolo- gist William Loomis ofthe University of California, San Diego, estimated in 1988 that “the important evolutionary differ- ences between a guppy and a primate probably lie in only a few hundred genes.” Along with the genetic contina- uum, and evolutionary continuities in structure, function and development, then, itis not surprising that one can dis cem a behavioral continuum. ‘Writing in 1980, the late zoologist John Paul Scott of Bowling Green State University pointed out that it isthe na- ‘ure ofthe environment that creates cer fain functional requirements for all or- ganisms if they are to survive. Like Rice's alga, a higher organism must take in nourishment and eliminate waste products. It must distinguish between predator and prey and between a poten- tial mate and a potential enemy. I must explore its environment and orient its sense organs appropriately as it takes in information about the beneficial and harmful aspects of its immediate world. (Organisms that ae relatively helpless at birth must have ways of indicating the need for care and nurturance. Only a few classes of adaptive behav- ior, Scott noted, are found in most species and at most phylogenetic levels. These include eating, the ight-or-fght response, sex, caregiving and investiga- tion. These patterns might be consid ered prototype adaptations. The con- nections between behavior and inner states and processes are less obvious. Yet such connections can be made by in- ference from a variety of evidence. This evidence includes knowledge of stir lating conditions, the effects of behav- {oral acts, knowledge of typical behav- ior patterns of the individual and species, choices made when alternatives, exist and reactions of other members of ‘one's group or species. A single overt display of emotions can reflect complex slates such as approach and avoidance, attack and fight, sex and aggression or fear and pleasure. It is not necessarily easy to detect an emotional substrate in the behavior of lower animals, but ne ther is itnecessary to exclude the possi bility. Emotion is far more complex than the subjective experience familiar to a Ihuman adult, and the concept of emo- tion can be applied to lewer animals as well as human beings. Emotions have an inherent complexity that isin part re- lated to their evolutionary history Cognition, Emotion and Evolution Any organism must determine, on the ‘basis of limited information, whether ‘there is food, a mate or danger in its en- vironment. Depending on the predic tion made, the organism makes a deci- sion to escape, to attack, to eat or to mate, The complex processes that go on in the service of biological need include receiving sensory input, evaluating it, capturing the important aspects of the information in symbols and comparing, the new information with memory stores, Predicting the characteristics of (=z | environments enables organisms to prepare for those environments. Psychologist Ulric Neisser of Comell University compared human beings with computing machines in a seminal article in 1963. Fle suggested that cogni- tive functions serve emotions and bio- logical needs. Information from the en- ‘vironment, he says is evaluated in terms ofits ability to satisfy or frustrate needs. What is particularly significant is that each new cognitive experience that is bi- logically important is connected with ‘an emotional reaction such as fea, plea- sure, pain, disgust or depression. From ‘the point of view of evolution, cognition developed in order to predict the future more effectively. The human brain, which has evolved as an adaptation 10 changing and difficult environments, has now helped create the very environ ‘met to which it must continue to adapt. If emotion is a chain of events, cogni- tion is generally near the beginning of the chain. This is considered an impor- tant point in the psychological commu- nity, which has put a good deal of effort [ saee [se Tmnpales oo into answering the “what comes first?” ‘question, ever since the American psy- chologist-philosopher William James in 1884 framed the question this way: Is it the feeling of emotion or the physiolog- ical changes that are part of emotion? This is actually a pseudoproblem. Emotions are not simply linear events, but rather are feedback processes. The function of emotion is to restore the in- dividual toa state of equilibrium when unexpected or unusual events create disequilibrium. Even if cognitions are generally at the beginning of the chain of events, they can be influenced by events appearing later in the chain— states of arousal, say, or ego defenses- through a feedback process. Stimulus events, either external or internal (as in dreams), act as primary triggers that start the emotion process going. ‘The biological aspects ofthis process have been the subject of considerable recent study. Animal research by Joseph E. LeDoux of New York Univ sity has revealed that the conditioned fear response involves several neural [over boravier] (me }- (ae) atest SS ae = oe < < < < Figure 4. Feedback loops in emotion show how sensory information is evaluated and translated into action or some other outcome that nor- _malizes the relationship between the individual and the triggering event. The inner tate perceived as fear may arise from a threat that is per~ Ceived as “danger; the feat triggers an impulse to flee, which results eventually in reduction of the threat, A similar set of homeostatic Processes can be seen inthe case of sadness ina child experiencing loss of her mother 2001 July-August 347 Figure 5. Although emotional substrates cannot always be discemed in the behavior of non- ‘human animals, many stimuli are experienced by people and animals alike and result pro- totypical behavior followed by, generally, the reesta might not have been achieved without the impulse pr ichment of an equilibrium state that tated by the inner state. In human ‘experience its common to use the term “emotion” to descube the feeling state, but in fact ‘emotion is considerably more complex. pathways with different latencies. ‘Damasio has traced the events in an ini- tial defensive response (Fea): The key features of a dangerous animal or event—perhaps color, speed of move- ‘ment, certain sounds—are detected and signaled to the amygdala, a part of the limbic system deep in the brain. This process is very rapid and is not a con- scious one. Signals from the amygdala to prefrontal areas and other locations precipitate the conscious felings ass0- ciated with an emotion. Feeling slates tend to be followed by impulses to action. Emotion can cause cones muscles to tense; it can be ex- pressed as a facial gesture, clenched fst ran action such as running, attacking or yelling. Impulses to action are not always Followed by action, as clinicians know— often fr fear of retaliation or embarrass- rent. Even when they are, overt behav ior isnot he end of the emotion process. ‘Such behavior generally has an effect on the stimulus or condition that started the chain of events in the first place. For example, running from a source of threat reduces the threat and tends to reestablish the condition that existed be- fore the threat. Similarly, f someone los- 5 parent crying and grieving tend to elicit supportive and helpful contacts from members of the grieving person's ‘348. American Scientist, Volume 89 social group and, at least in a symbolic ‘way, provide a sort ofreattachment with the lost parent andl thus a change in the feeling state. Overall, emotion isa kind of homeo static process in which behavior medi- ates progress toward equilibrium; call it a behavioral homeostatic, negative- feedback system. Emotion isa chain of events made up of feedback loops. Feel ings and behavior can affect cognition, just as cognition can influence feeling ‘At the heart of all these descriptions is theidea that emotions have a function in the lives of individuals, This idea arises from an evolutionary perspective, is can- sistent with psychodynamic thinking and isbecoming increasingly acepted in con- temporary writings. For example, young organisms require food, protection ancl ‘transportation. Crying is amajor method for getting such care. Fear protects the self initiates withdrawal and allows gen- eral functioning to continue. Shame leads to remorse and a decrease in the probe- bility ofrepeition ofthe shameful act. These examples imply that emotions are patt of a social regulation process. Evolutionary theory reminds us that the interests of diferent individuals are of- ten in conflict males versus females, pparents versus children brothers versus brothers, group versus group. Genes are “selfish and concemed with selfmain- tenance and self reproduction. Social ir- teractions and communications reflect this conflict. Listening and speaking are regulated by direct and subtle expres- sions of emotion—smiles, eye contact or looking away, nods, postural shifts, vo- calizations, passive questions and im- plicit commands (“Why don’t you wear a hearing aid?”) ‘Much social interaction involves in- dividuals in different hierarchical po- sitions. In such situations, during con- versation there is often a censoring of rebellions thoughts, and a covert compe tition of ideas. An individual may expe- rience felings of defiance or may accept a submissive position. Emotions, at both conscious and unconscious levels, regu late such socal processes. Another way to conceptualize emo- ‘tions as a social-regulation process is in terms ofthe views of animal communi- cation proposed by Eugene Morton of the Smithsonian Institution and his col- Teagues. They point out that communi cation is an assessment/management process aimed at survival, Communica- tion signals are selected in evolution be- cause they substitute for more risky be- havior suchas fighting. California ground squirrels stimulate a rattlesnake lo rattle by kicking earth at it. Thisis done because squimels use the sound of rattling to assess the snake's size and body temperature, two factors that determine how dangerous the snake is to their pups, With many ant mals, distress cals are adaptive because they may startle a predator into letting 0, may attract the attention of other Conspecifics to mob the predator, or may attract a larger predator to compete and possibly allow escape, However, itis not always easy to determine the adaptive nature ofa given signal Emotions are part of the management of the process. Anges, for example, in- timidates, influences others to do some- thing you wish them to do, energizes an individual for attack or defense and spaces the participants in a conflict. Modeling the Emotions have used the term “emotion” asa sin- gle, general tem for a group of phenom- ena. As complex processes with func- tional value both in communication and in increasing the individual's chances of survival, emotions represent proximate methods to achieve evolutionary fitness. To integrate many of the things known about emotions, model makings useful. In English there are a few hundred emotion words, and they tend to fll into families based on similarity. have found. that the primary emotions can be con- ina fashion analogous to a color wheel—placing similar emotions close together and opposites 180 degrees apart, like complementary colors. Other emotions are mixtures of the primary emotions, just as some colors are prima- sy and others made by mixing the pri- sary colors. Such “circumplex’ model- ing can be used as an analytical tool in understanding personality 25 well, and the similarity between the two models is, important. [have extended the circum- plex model into a third dimension, rep- resenting the intensity of emotions, 50 that the total so-called structural model of emotions is shaped like a cone. The notion of a circumplex mode! is ‘not my invention, nor is it new. Social ‘psychologist William McDougall noted the parallel between emotions and colors in 1921, wnting that “the color sensations present, lke the emotions, an indefinitely great variety of qualities shading into one another by imperceptible gradiens...” ‘The first creumplex model was one de- veloped by Brown University psycholo- gist Harold Schlosberg in 1941, after he hnad asked research participants to judge the emotions posed in a standard set of pictures of facial expression. Schlosberg, added the intensity dimension to his model. My own model was proposed in. 1958, when [suggested eight basic bipo- lar emotions: joy versus sorrew, anger Ver sus far accepance versus disgust and sir~ prise versus expectaricy. Over the centuries, from Descartes to the present, philosophers and psycholo- gists have proposed anywhere from 3 to i emotions as primary or basic. All the lists include far, anger and sadness; most include jay, love and surprise. There is ro unequivocal way to seitle on a precise number, although factor-analytic stud- ies, similarty-scaling studies, child-de- velopment studies and cross-cultural studies are useful. But in the final analy- sis, this is a theoretical decision to be Figure 7. To help psychologists understand the relations among emotions, raters were asked to estimate the degree of similarity be tween certain pais of emotions. The use ofa Smilaity sealing method produced the em pirical angular locations shoven in the figure (where opposites havea similarity ranking of LO and identical concepts have a similarity ranking of L.0) The concepts shown are 2 e- lection from a larger population of words, Figure 6, Author's threeimensional crcumplex model describes the relations among emotion ‘concepts, which are analogous t the colors on color wheel. Te cones vertical dimension rep- ‘eoents intensity, and the cicle represent degrees of smarty among the emotions. The eight ectors are designed to indicate that there ae eght primary emotion dimensions defined by the theory arranged as four pais af opposites. In the exploded model the emotions in the blank spaces aze the primary dyads-emotions that are mixtures of two ofthe primary emotions. accepting ful fe 7 7 a 3 2001 July-August 348 evaluated in terms ofthe inferences and {insights to which it leads, the research it suggests and the extent to which empir- ical data are consistent with it The psy- choevolutionary theory assumes there are eight basic emotion dimensions arranged in four pairs. If there are eight basic emotion dimen- sions (each with a number of synonyms or related terms), how can we account forthe total language of emotions? Vari- cous published studies imply that the few ‘huncired emotion words tend to fall into farnilies based on similarity. we follow the pattern used in color theory and re- search, we can obtain judgments about combinations—the emotions that result ‘when two or more fundamental emo- tions are combined in the same way that red and blue make purple. Judges in these studies have agreed that mixing joy and acceptance produces the mixed emo- tion of love; disgust plus anger produces hatred or hostility. Such mixtures have been called primary dyats in the theory. One can continue on this way and ac- count for hundreds of emotion terms by mixing two or more emotions at diffes- ent levels of intensity ‘As noted above, itis interesting and ‘pethaps important that one of the hy- potheses generated from this structural model is that personality traits should have a similar structure. Again we can take cues from language. Although per- sonality is usually taught in universities as if it had little or nothing to do with ‘emotions, words such as gloomy, resent- ful, anxious and ci can describe person- ality traits as well as emotional states. An individual can fel depressed, orbea de- pressed person, feel nervous or be aner- ‘vous person. Often people are able to ‘measure both emotional states and per- sonality traits using the same checklist of adjectives, with a simple change in in- structions. When research participants are asked how they feel now, or within the past few days, the instruction asks fora self-report ofan emotional state ora ‘mood. But they can be asked how they usually feel, a question that yields infor- ration about personality traits. At the extremes are pathological states such as mania and paranoia—but even these can, bbe conceived as extreme expressions of such basic emotions as sadness, joy and disgust, Thus personality traits may be conceptualized as being derived from mixtures of emotions, With my colleague ‘Hope Conte, I have been able to find a circumplex structure for certain classes ofpersonality traits. 350. American Scientist, Volume 89 Furthermore, in recent years there have been more than 100 published studies concemed with identifying per- sonality characteristics in lower ani- ‘mals, Of course lower animals probably should be said to have temperament rather than “personality,” but extraver- sion, emotional stability and agreeable- ness have shown considerable generali- ty across species. Samuel Gosling of the University of Texas at Austin and Oliver John of the University of California, Berkeley, have identified extraversion and emotional stability as characteris- fics of animals as low on the phyloge- netic scale as guppies and octopuses. Some Implications ‘An evolutionary framework supplies the study of emotions with such con- cepts as functional thinking, the gener- ality of mechanisms across phyla, de- velopmental theory (to explain, for instance, attachment), and the concepts of inclusive fitness and proximate and, ultimate causation. Happily, in combination with the se-