The Behavioral Ecology View of Facial Displays, 25 Years Later


Alan Fridlund

Alan J. Fridlund, Department of Psychological and Brain Sciences, UC Santa Barbara

Copyright © by Alan J. Fridlund, All Rights Reserved.

The current version benefitted from the editorial efforts of Andrea Scarantino. An expanded version will appear in a volume published by Oxford University Press.

August 2015 – I began as an adherent of the Basic Emotions Theory (BET) of facial expressions. In most formulations, BET held that emotions, understood as internal states or discrete affect categories, were associated with specific patterned movements termed “facial expressions of emotion.” The foundation for BET was research in which members of diverse cultures matched a small number of photos of posed facial expressions to a similarly small number of emotion terms, suitably translated or mapped onto stories (Ekman, 1972; Ekman & Friesen, 1971; Ekman, Sorenson & Friesen, 1969; Izard, 1971).

These matching-to-sample studies, jointly with other evidence, were declared to mean that: (1) the emotions signified by the terms/stories were “biologically based,” i.e., they were phylogenetic; (2) the emotional facial expressions matched to the emotion terms were uniform in their production and universal in their recognition; and (3) there was an automatic, causal link between the prototypical emotional faces and the respective internal emotional mechanisms (collectively, the “Facial Affect Program”) that produced them. In BET, any deviation from the predicted correspondence between a triggered emotion and the emission of its counterpart facial expression was due to the intervention of cultural “display rules” governing social behavior (Ekman & Friesen, 1969). Such culturally dependent control was imperfect, however, and so a muted, throttled or distorted expression might “leak” traces of the suppressed, genuine emotion of the expressor onto the face.

At the start of my career, BET was the dominant framework for studying emotion and facial expression, and I had no reason to challenge it. I began by conducting electromyographic studies of the tiny facial movements people made during emotional imagery (Fridlund, Schwartz & Fowler, 1984), work begun by Paul Fair and Gary Schwartz (Schwartz, Fair et al., 1976). Gary invited me to his Yale lab to conduct my doctoral studies, and introduced me to Silvan Tomkins. Later he arranged for me to meet Carroll Izard and Paul Ekman, the two leading BET theorists at the time. I came to know both men well, and I may be the only person to have written papers with each (Ekman & Fridlund, 1987; Fridlund, Ekman, & Oster, 1988; Fridlund & Izard, 1983; Matsumoto, Ekman, & Fridlund, 1990). Over time, however, I developed unresolvable disagreements with them over the tenets of BET.

My skepticism about BET came from four growing realizations: (1) the cross-cultural findings could never have been helpful in apportioning roles to “biology” vs. “culture,” because both diversity and uniformity can arise from natural selection (think finches); (2) claiming cross-cultural uniformity for certain iconic facial expressions after obtaining matches to emotion categories, and universality of those “basic emotions” based on the same matches, was circular and tautological; (3) on closer inspection, the matching between facial displays and emotion terms/stories began to appear inflated to me and other researchers, due to technical deficiencies in the experimental protocols; and, most important, (4) regarding the face as an automatic but suppressible readout of internal, “authentic” emotional states conflicted with modern views of animal communication. This last point was most critical in convincing me that BET was fatally flawed.

Darwin first attempted systematically to link animal signaling with our facial expressions, and BET theorists duly pay homage to him, but misread him when they cite him to support their claim that our facial expressions evolved “to express emotion” (see Fridlund, 1992a). In promulgating evolution by natural selection, Darwin had first to dispose of a contending position: the Argument from Design, made by Charles Bell, William Paley and others, which held that creatures were well-suited to their niches because God made them so. Thus, in On the Origin of Species, Darwin could not repeat identical evidence of goodness-of-fit and then argue a different conclusion. Instead, he used evidence of imperfect design – vestigial structures such as webbed feet on land birds, phalanges in a seal’s flipper, and the human appendix – as proof of common origins and to vitiate notions of perfect design ex nihilo (Browne, 1985; Darwin, 1859; Fridlund, 1992a; Gruber, 1974).

In The Expression of the Emotions in Man and Animals, Darwin extended his assault on the Design argument to Bell’s view of facial expressions: “I want, anyhow, to upset Sir C. Bell’s view … that certain muscles have been given to man solely that he may reveal to other men his feelings” (F. Darwin, 1887, Vol. 2, p. 78.). He proposed instead that, as with vestigial organs, most facial behaviors were likewise rudimentary and “of no service, often of much disservice,” or “purposeless” (Darwin, 1872, pp. 67, 76). They were either remnants of reflexes that had been useful (“serviceable associated habits”), antithetical remnants arising from contrasting elicitors (“antithesis”), or overflows of excitation (“direct actions of the nervous system”). Any communicative value for facial expressions was incidental. Although Darwin’s account neutered Bell’s creationism, it left him unequipped to argue that facial expressions evolved for anything (Fridlund, 1992a).

While honoring Darwin, BET actually co-opted the 1950’s mechanistic ethology of Lorenz and Tinbergen (Lorenz, 1967, 1970; Tinbergen, 1952, 1953). These early ethologists’ tripwire fixed-action patterns became the “Facial Affect Program” of the neurocultural version of BET (even the “FAP” acronyms were identical; Fridlund, 1992b). Instead of red spots on beaks that released appeasement displays and food calls, prototypical emotional events triggered the release (“expression”) of emotion that spilled out on our faces and reflected our “true feelings” except if modified by tradition (“display rules”), training, or treachery.

If BET was drawing upon the Lorenz-Tinbergen formulations, modern ethology was abandoning them, leaving BET gutted of its claim to Darwin’s imprimatur and severing BET from any continuity with developing models of animal communication. Animal behaviorists began to note that most nonhuman signals didn’t look fixed or cartooney, but flexible, social and contextual (Alcock, 1984; Hinde, 1985a, b; Smith, 1977). Such behavioral ecologists (cf., Davies, Krebs, & West, 2012; Maynard Smith, 1982) saw animal signaling not as vestigial reflexes, or readouts of internal state, but as adaptations that served the interests of signalers within their social environments. Signaler and recipient – even when predators and prey (e.g., “pursuit deterrence” signals; see Caro, 2005) – were reconceived as coevolved dyads in which displays indicated the likely behavior of issuers, with recipients using such behavior as cues to the issuers’ next moves (Krebs & Davies, 1987; Krebs & Dawkins, 1984).

Although Darwin’s vestigial reflexology in Expression was outdated, it seemed to me that modern behavioral ecology’s view of expressive behavior as dynamic and contextual offered a way to resurrect Darwin’s grander vision of continuity between human and nonhuman signaling. Thus, in the 1990’s, I began writing position papers and conducting studies on what became the Behavioral Ecology View (BECV) of human facial behavior. In this account, human facial displays, like animal signals, serve the momentary “intent” of the displayer toward others in social interaction (Fridlund, 1990; 1991a, b; 1992a, b; 1994; 1996; 2002; 2006). (“Intent” here is adduced from people’s interactional trajectory; it does not presuppose that people know, can articulate, and/or will disclose what they intend).

Indeed, many of the classic, iconic BET expressions can be recast in such intentional, functional terms. In suitable contexts, so-called “happy faces” solicit affiliation or play, whereas “sad faces” recruit succor, “anger faces” threaten or deter, “fear faces” predict submission or withdrawal, “disgust faces” indicate rejection or intent to spew, and so on (see Table 1 below).

table alan

Table 1: Two Views of Facial Expressive Behavior: BET’s “Facial Expressions of Emotions” vs. BECV’s Functional Social Tools


When participants are asked to match iconic BET expressions to functional redescriptions (e.g., “back off or I’ll attack”), such redescriptions achieve matching rates equal to emotion terms (e.g. “anger”; Yik, 1999). Unlike emotion terms, however, these functional descriptors imply neither any particular internal state (e.g. one can solicit affiliation or play when happy or unhappy), nor any moral assignations about which signals are “honest” or “genuine” (e.g. a face recruiting succor is not “honest” because one is sad and dishonest when one is not). Such functional descriptions are predicated merely on the view that facial displays are only probabilistic signals of social intentions that would, in everyday life, be accompanied by the words, vocal prosodies, and gestures congruent with the intent.

According to BECV, facial displays serve as social tools. To wit, in accusing a relationship partner of committing an infidelity, one might exclaim, “You are a stinking, lying turd!” and supply the concordant tone of voice, an upturned nose, and the appropriate hand and finger gestures. All this sound and fury would force a nixing or resetting of the relationship. For BECV, understanding that the shock-and-awe display was a tool for relationship realignment is all it takes to explain why the signaling occurred. Any detour to qualia (or other internal proxy for emotion) as causal is extraneous because, in BECV, there is no necessary connection between those signals and any one emotion: the accuser/displayer may have been disgusted, contemptuous, devastated, livid – or relieved or thrilled, if the entire rejection drama was staged to divert the partner from discovering that he/she cheated first.

BET advocates objected to this interactional view of facial displays, noting that “Facial expressions do occur when people are alone … and contradict the theoretical proposals of those who view expressions solely as social signals” (Ekman, Davidson & Friesen, 1990, p. 351). It was necessary, therefore, to explain that being alone physically didn’t imply that we were alone psychologically.

In this social media age, when people have their faces glued to their phones and begin and end relationships with right and left swipes, this explanation now seems obvious, but it was originally contentious. From the start, I had enumerated examples in which we were alone but implicitly social (Fridlund, 1991a): imagining or misbelieving that others are present (daydreams, flashbacks, or talking to someone who’s left the room), grieving (when we crave reunion), sexual fantasy, soliciting an interaction (recruiting succor with a pained or crying face, as infants do), or preparing for one (rehearsing for a play or interview).

In all these cases, individuals may subvocalize – they are “talking to people in their heads” – and any accompanying “solitary” faces would be equally social. It makes no difference if the interactant is myself: If I scowl and tell myself, “Now Fridlund, don’t screw up!”, both my words (sotto voce) and accompanying face (sotto facie?) serve to keep Fridlund focused and out of trouble.

We showed this experimentally, with human studies that extended novel avian research by the much-missed Peter Marler (Marler, Duffy & Pickert, 1986a, b). We demonstrated audience effects in solitary smiling with audiences that were both explicit (friends were present) and implicit (participants were alone but believed friends were co-participants elsewhere), and with social vs. nonsocial imagery (Fridlund 1991, Fridlund et al., 1990, 1992). Several investigators have replicated such implicit audience effects, expanding the findings to infants, beyond smiling, and to augmenting vs. decrementing effects of friends vs. strangers (Hess, Banse & Kappas, 1995; Jones, Collins & Hong, 1991; Schützwohl & Reisenzein, 2012; Wagner & Smith, 1991).

BET partisans dismissed these findings peremptorily: “No account should be taken of studies that do not distinguish between Duchenne and non-Duchenne smiles…” (Ekman & Keltner, 1997, p. 41). “Duchenne smiles,” according to BET, were genuine, emotional, “felt” smiles, unlike other, intrinsically social smiles, which might be “false,” “phony” or “unfelt” (Ekman & Friesen, 1982; Frank & Ekman, 1993; Frank, Ekman & Friesen, 1993). The criticism was entirely misplaced, since in the implicit-audience studies, the smiles in question varied substantially with sociality but were all emitted in solitude – which, for BET, would make them emotional and genuine (again, Ekman et al., 1990, p. 351).

As Ruth Leys noted (personal communication, March 5, 2015), Ekman soon changed course from dismissing these implicit-sociality findings to revising his theory to accommodate them: “I expect that some display rules are so well established that some people may follow them even when they are alone. And some people when alone may imagine the reactions of others, and then follow the appropriate display rule, as if the others were present. And finally, there may be display rules that specify the management of expression not just with others but when alone” (Ekman, 1997, p. 328). Notably, Ekman did not specify how one might ascertain when such “solitary display rules” were in effect and when they were not.

If Ekman’s turnabout solved one problem, it opened up a bigger one. Prior to this change, Ekman contended that solitary facial behavior was free of display rules. Of the paradigmatic Japanese-American study cited most as a demonstration of the display-rules concept (Ekman, 1972; Friesen, 1972), Ekman summarized the findings: “In private, when no display rules to mask expression were operative, we saw the biologically based, evolved, universal facial expressions of emotion. In a social situation, we had shown how rules for the management of expression led to culturally different facial expressions” (Ekman, 1984, p. 321). With Ekman’s expansion of BET to include solitary display rules, can it now be certain that the solitary faces observed in the Japanese-American study were display-rule-free and thus “biologically based, evolved, universal facial expressions of emotion”? If so, how would that be verified?

There are wider repercussions. Ekman’s concession that private behavior may be conventional like our public behavior reduces considerably the distance between the claims posed by his neurocultural version of BET, and those struck earlier by the cultural relativists he so staunchly opposed, such as Margaret Mead and Ray Birdwhistell, who argued for the pervasiveness of cultural learning in all aspects of life.

Findings that solitary smiles could be “social” also seemed to violate BET’s felt/false, Duchenne/non-Duchenne smile dichotomy. Can this dichotomy be sustained? I proposed early on that “Duchenne” smiles were actually conjoint displays of typical smiling plus tonic elicitation of the blink reflex of Descartes (“wincing”), the latter of which could occur with any strong stimulus and not any specified emotional state (Fridlund, 1994).

Studies now indicate that, contrary to BET, Duchenne smiles are at least as affected by sociality as non-Duchenne ones (Crivelli, Carrera & Fernández-Dols, 2015; Fernández-Dols & Ruiz-Belda, 1995; Mehu, Grammer & Dunbar, 2007; Ruiz-Belda, Fernández-Dols & Barchard, 2003), that they can be produced deliberately (Gosselin, Perron & Beaupré, 2010; Gunnery & Hall, 2014; Gunnery, Hall & Ruben, 2013), and that their occurrence varies both with smile intensity (Krumhuber & Manstead, 2009) and stimulus intensity regardless of valence (Harris & Alvarado, 2005).

BECV rejects the idea that some display, or class of displays, can have intrinsic properties outside the context of its issuance. Smiles may be made by mothers toward children or assailants toward victims. Tears may flow in grief, retribution, reconciliation, or triumph. The meanings of these displays can be understood only by considering who makes them and when they occur.

In the neurocultural version of BET, however, morphology dictates not just emotionality but authenticity. Duchenne smiles are “genuine” because they are “felt,” and disingenuous because they are “unfelt” or “false.” This stark dichotomy turns everyday courtesy into mendacity. It also leads to futile diversions. A used-car salesman may be a consummate Duchenne smiler and scam nearly every customer who walks onto his lot. His winning Duchenne smiles sell cars. For BET, then, his smiles must be “felt.” Is this what we care about, whether he’s happy if he scams us?

For BECV, the “authenticity” of his smile lies not in what he feels, but in whether it predicts whether he will treat us fairly if we buy a car from him. More generally, we learn whose words and expressions are reliable indicators of their intent, and over time we bond with those individuals who prove reliable and avoid those who prove otherwise.

In deception, therefore, the “truth” of a display inheres neither in the display nor its displayer, but in the moving average by which a recipient continually calibrates and recalibrates the reliability of the signals issued in that context by that displayer. Greater predictability of displayers’ signals and lower skepticism by recipients toward those signals naturally co-evolve with repeated cooperation, else breaches occur that force recalibration, confrontation, or termination of interaction (Mitchell & Thompson, 1986). The “leakage” seen by BET theorists as the breakout of “genuine emotion” through an outer mask is simply a momentary conflict in intentions in social negotiation (Fridlund, 1991a). This interactional perspective is decidedly anti-Darwin qua Expression but resoundingly Darwinian (Fridlund, 1992a).

Certain questions have been raised repeatedly about BECV. Does BECV deny “emotion”? Does BECV deny a privileged relationship between “emotion” and certain facial displays? Do “emotions” serve as “commitment devices” that reveal our authentic, internal states (Frank, 1988; and see crucial treatment of the emotion-as-commitment issue by Leys, 2013)? To BECV, all these questions mean little, because they hinge entirely on how one defines emotion (cf., Schattschneider, 1960; to paraphrase: defining the terms determines the outcome). Over a century’s theory and research have demonstrated that “emotion” has proven intractable to consensual, let alone operational, definition.

BET theorists often identify “emotion,” at least implicitly, with qualia or “feelings.” For example, an observer may claim that someone “felt sad” and his sadness produced his “sad expression.” In the neurocultural version of BET, “felt” (“Duchenne”) smiles are “all smiles in which the person actually experiences … a positive emotion” (Ekman & Friesen, 1982, p. 242). Both the “sadness” and “positive emotion” contentions make qualia – or their putative proximal generators – causal, and both are untenable when they make accountable something ineffable and unverifiable.

But what if one were to localize the proximal generators for qualia, the “feeling centers,” in the brain? Could we then say that changes in qualia, or those generators that produced the qualia changes, caused the associated events in the neuromuscular centers that produced the facial expressions? How does one ever determine that event A causes event B in the brain? I invite readers new to this question to Google “Libet’s experiment” and discover the labyrinthine complexities in determining neurocausality.

One common BET recourse to according qualia strict agency and keep “emotion” scientific is to make facial expressions just part of the package of changes (neurochemical, behavioral, cognitive) that constitutes an emotion or “affect program,” qualia being among them. On this view, the presence or absence of emotion cannot be determined by the presence or absence of qualia, or of any other single component or subset of components.

This view reduces to no more than hand-waving about the knottiness of the phenomena and ad hoc choices of stipulated “emotion measures,” with the result that surveys of research and formal meta-analyses continually find disappointing links between “emotion” and “expression” (cf., Ortony & Turner, 1990). Newer backstops include: (1) trying to re-objectify “emotion” as a neo-Kantian, categorical “conceptual act” that belongs more to the observer than the emoter (Barrett, Wilson-Mendenhall, & Barsalou, 2015), and (2) paradoxically trying to nail down the “emotion” concept by declaring it intrinsically fuzzy (Scarantino & Griffiths, 2011).

For BECV, all this reasoning is tendentious and wasteful if the purpose is to understand our facial displays. The same holds for ecumenical BET formulations that begin with emotion, variously defined, and end with how “everyone knows” that the expressions have social functions, too (e.g., Hauser, 1996). For BECV, displays evolved as social tools directly, not as parts of underlying mechanisms for the production of displays. Natural and cultural selection do not “care about” (specifically select for) the inner workings of traits, only the traits themselves.

Facial behaviors that aid individuals in navigating their social terrains (i.e., displays) will, via their displayers, tend to proliferate horizontally (i.e., culturally and geographically) and vertically (via genetic/epigenetic inheritance), regardless of what neural operations produce them; accompanying these displays is the coevolution of recipient behavior that is attentive yet skeptical (Krebs & Dawkins, 1984).

How has BECV fared against BET? James Russell’s influential critique of the cross-cultural matching-to-sample studies (Russell, 1994), and his team’s demonstration of powerful context effects in facial-expression perception (Carroll & Russell, 1996; Russell & Fehr, 1987), broke the paradigm lock BET had on facial-expression research. BECV’s small contribution has been to supply a new framework for understanding our facial displays, one that restores Darwin’s vision of human-animal continuity and places it on a solid evolutionary footing. I believe it’s what Darwin would have proposed had he been able.

I am pleased by how much serious scholarly attention BECV has received. I grounded it in behavioral ecology and evolutionary theory, but Brian Parkinson’s generous review reminded me of its debt to Dewey (Parkinson, 2005). With penetrating depth, Ruth Leys has shown how BECV can clarify philosophical and technical problems in the objectification and neural localization of emotion (Leys, 2007, 2010, 2011, 2014). BECV has informed research on both public and implicit-audience accounts of responses to social media (Litt, 2012), smiling in pain (Kunz, Prkachin, & Lautenbacher, 2013), human-computer communication (Aharoni & Fridlund, 2007), persuasion (Cesario & Higgins, 2008), power and dominance (Burgoon & Dunbar, 2006), facial displays in rats (Nakashima, Ukezono, Nishida, Sudo, & Takano, 2015) and chimpanzees (Parr & Waller, 2006), intrapersonal communication in therapeutic narrative writing (Brody & Park, 2004), and the game-theoretic analysis of human deception (Andrews, 2002).

Finally, José-Miguel Fernández-Dols and his colleagues have conducted a line of masterful studies showing how BECV can account for facial behavior in naturalistic settings (e.g., Crivelli et al., 2015; Fernández-Dols & Ruiz-Belda, 1995; Ruiz-Belda et al., 2003). It also seems that the battle royale between BET and BECV has liberated inquiry on facial expressions: investigators can now pursue hypotheses (e.g., genetic/epigenetic diversity in facial displays, facial dialects, infant deception) that, because they transgressed BET, were previously inconceivable or taboo.

BECV will always be a tough sell. It requires shaking off a romanticized view of human nature that makes the face a battleground between an “authentic self” and an impression-managed “social self” (Fridlund, 1994, 1996). The first concept we treasure; the second we concede reluctantly. To BECV, both are illusory. Like our words, voice and gestures, our facial displays – even those we make as infants, and which will be deployed by our android companions (will they have felt or false Duchenne smiles?) – are part of our plans of action in social commerce.


I am indebted to Jose-Miguel Fernández-Dols and Ruth Leys for comments and suggestions.


Aharoni, E., & Fridlund, A. (2007). Social reactions toward people vs. computers: How mere labels shape interactions. Computers in Human Behavior, 23, 2175-2189.

Alcock, J. (1984). Animal behavior: An evolutionary approach (3rd. Ed.). Sunderland, MA: Sinauer.

Andrews, P. W. (2002). The influence of postreliance detection on the deceptive efficacy of dishonest signals of intent: Understanding facial clues to deceit as the outcome of signaling tradeoffs. Evolution and Human Behavior, 23, 103-121.

Barrett, L. F., Wilson-Mendenhall, C. D., & Barsalou, L. W. (2014). The conceptual act theory: A road map. In L. F. Barrett and J. A. Russell (Eds.), The psychological construction of emotion (pp.83-110) New York: Guilford.

Block, N. J., Flanagan, O. J., & Güzeldere, G. (1997). The nature of consciousness: Philosophical debates. Cambridge: MIT Press.

Brody, L. R., & Park, S. H. (2004). Narratives, mindfulness, and the implicit audience. Clinical Psychology: Science and Practice, 11, 147-154.

Browne, J. (1985), Darwin and the expression of the emotions. In D. Kohn (Ed.), The Darwinian heritage (pp. 307-326). Princeton: Princeton University Press.

Burgoon, J. K., & Dunbar, N. E. (2006). Nonverbal expressions of dominance and power in human relationships. Sage handbook of nonverbal communication (pp. 279-297). Thousand Oaks: Sage.

Caro, T. M. (2005). Antipredator defenses in birds and mammals. Chicago: University of Chicago Press.

Carroll, J. M. & Russell, J. A. (1996). Do facial expressions express specific emotions? Judging emotion from the face in context. Journal of Personality and Social Psychology, 70, 205-218.

Cesario, J., & Higgins, E. T. (2008). Making message recipients “feel right”: How nonverbal cues can increase persuasion. Psychological Science, 19, 415-420.

Chalmers, D. (1995). Facing Up to the Problem of Consciousness. Journal of Consciousness Studies, 2, 200–219.

Crivelli, C., Carrera, P., & Fernández-Dols, J.M. (2015). Are smiles a sign of happiness? Spontaneous expressions of judo winners, Evolution and Human Behavior, 36, 52-58.

Darwin, C. R. (1859). On the origin of species, or the preservation of favoured races in the struggle for life. London: Murray.

Darwin, C.R. (1872). The Expression of the Emotions in Man and Animals. London: Murray.

Darwin, F. (Ed.) (1887). The life and letters of Charles Darwin, including an autobiographical chapter. (Vols. 1 and 2). London: Murray.

Davies, N. B., Krebs, J. R., & West, S. A. (2012). An introduction to behavioral ecology (4th Ed.). Oxford: Wiley-Blackwell.

Ekman, P. (1972). Universals and cultural differences in facial expressions of emotion. In J. Cole (Ed.), Nebraska Symposium on Motivation, 1971 (Vol. 19), (pp. 207-282). Lincoln, NE: University of Nebraska Press.

Ekman, P. (1997). Expression or communication about emotion. In N. L. Segal, G. E. Weisfeld, & C. C. Weisfeld (Eds.) Uniting Biology and Psychology: Integrated Perspectives on Human Development (Pp. 315-338). Washington, D.C.: APA Press.

Ekman, P., Davidson, R. J., & Friesen, W. V. (1990), The Duchenne smile: Emotional expression and brain physiology II. Journal of Personality and Social Psychology, 58, 342-353.

Ekman, P., & Fridlund, A. J. (1987). Assessment of facial behavior in affective disorders. In J. D. Maser (Ed.), Depression and expressive behavior. Hillsdale, New Jersey: Erlbaum.

Ekman, P. & Friesen, W. V. (1969). The repertoire of nonverbal behavior: Categories, origins, usage, and coding. Semiotica, 1, 49-98.

Ekman, P., & Friesen, W. V. (1971). Constants across cultures in the face and emotion. Journal of Personality and Social Psychology, 17, 124-129.

Ekman, P., & Friesen, W. V. (1982). Felt, false, and miserable smiles. Journal of Nonverbal Behavior, 6, 238–258.

Ekman, P., & Keltner, D. (1997). Universal facial expressions of emotion: An old controversy and new findings. In U. Segerstrâle & P. Molnár (Eds.), Nonverbal communication: Where nature meets culture (pp. 27-46). Mahway, N. J.: Erlbaum.

Ekman, P., Sorenson, E. R., & Friesen, W. V. (1969). Pan-cultural elements in facial displays of emotions. Science, 164, 86-88.

Fernández-Dols, J.M., & Ruiz-Belda, M.A. (1995). Are smiles a sign of happiness? Gold medal winners at the Olympic Games. Journal of Personality and Social Psychology, 1995, 69, 1113-1119.

Frank, M. G., & Ekman, P. (1993). Not all smiles are created equal: The differences between enjoyment and nonenjoyment smiles. Humor: International Journal of Humor Research, 6, 9-26.

Frank, M. G., Ekman, P., & Friesen, W. V. (1993). Behavioral markers and recognizability of the smile of enjoyment. Journal of Personality and Social Psychology, 64, 83–93.

Frank, R. H. (1988). Passions within reason: The strategic role of the emotions. New York: Norton.

Fridlund, A. J. (1991a). Evolution and facial action in reflex, social motive, and paralanguage. Biological Psychology, 32, 3-100.

Fridlund, A. J. (1991b). The sociality of solitary smiles: Effects of an implicit audience. Journal of Personality and Social Psychology, 60, 229-240.

Fridlund, A. J. (1992a), Darwin’s Anti-Darwinism in The Expression of the Emotions in Man and Animals. In K. Strongman (Ed.), International review of emotion, Vol. 2.

Fridlund, A. J. (1992b). The behavioral ecology and sociality of human faces. In M. S. Clark (Ed.), Review of Personality and Social Psychology, 13, 90-121. Newbury Park, CA: Sage.

Fridlund, A. J. (1994). Human facial expression: An evolutionary view. San Diego, CA: Academic Press.

Fridlund, A. J. (1997). The new ethology of human facial expressions. In J. A. Russell & J.-M. Fernandez-Dols (Eds.), The psychology of facial expression (pp. 103-129). Cambridge: Cambridge University Press.

Fridlund, A. J. (2002). The behavioral ecology view of smiling and other facial expressions. In M. Abel (Ed.), An empirical reflection on the smile (pp. 45-82). New York: Edwin Mellen Press.

Fridlund, A. J., Ekman, P., & Oster, H. (1988). Emotions and facial expressions. In A. Kendon (Ed.), International encyclopedia of communications. Philadelphia: Annenberg School of Communications/Oxford University Press.

Fridlund, A. J., & Duchaine, B. (1996). ‘Facial Expressions of Emotion’ and the delusion of the hermetic self. In R. Harré & W. G. Parrott, The emotions (pp. 259-284). Cambridge: Cambridge University Press.

Fridlund, A. J., & Izard, C. E. (1983). Electromyographic studies of facial expressions of emotions and patterns of emotions. In J. T. Cacioppo & R. E. Petty (Eds.), Social psychophysiology: A sourcebook (pp. 243-286). New York: Guilford Press.

Fridlund, A. J., Kenworthy, K., & Jaffey, A. K. (1992). Audience effects in affective imagery: Replication and extension to dysphoric imagery. Journal of Nonverbal Behavior, 16, 191-212.

Fridlund, A. J., & Russell, J. A. (2006). The functions of facial expression: What’s in a Face? Sage handbook of nonverbal communication (pp. 299-319). Thousand Oaks: Sage.

Fridlund, A. J., Sabini, J. P., Hedlund, L. E., Schaut, J. A., Shenker, J. I., & Knauer, M. J. (1990). Social determinants of facial expressions during affective imagery: Displaying to the people in your head. Journal of Nonverbal Behavior, 14, 113-137.

Fridlund, A. J., Schwartz, G. E., & Fowler, S. C. (1984). Pattern-recognition of self-reported emotional state from multiple-site facial EMG activity during affective imagery. Psychophysiology, 21, 622-636.

Gosselin, P., Perron, M., & Beaupré, M. (2010). The voluntary control of facial action units in adults. Emotion, 10, 266–271.

Gruber, H. E., with Barnett, H. P. (1974). Darwin on man: A psychological study of scientific creativity. New York: Dutton.

Gunnery, S. D., & Hall, J. (2014). The Duchenne smile and persuasion. Journal of Nonverbal Behavior, 38, 181-194.

Gunnery, S. D., Hall, J. A., & Ruben, M. A. (2013). The deliberate Duchenne smile: Individual differences in expressive control. Journal of Nonverbal Behavior, 37, 29-41.

Harris, C. R., & Alvarado, N. (2005). Facial expressions, smile types, and self-report during humor, tickle, and pain. Cognition and Emotion, 19, 655-669.

Hauser, M. (1996). The evolution of communication. Cambridge: MIT Press.

Hess, U., Banse, R., & Kappas, A. (1995). The intensity of facial expression is determined by underlying affective state and social situation. Journal of Personality and Social Psychology, 69, 280-288.

Hinde, R. A. (1985a). Expression and negotiation. In G. Zivin (Ed.), The development of expressive behavior (pp. 103-116). Orlando, FL: Academic Press.

Hinde, R. A. (1985b). Was ‘The Expression of the Emotions’ a misleading phrase? Animal Behaviour, 33, 985-992.

Izard, C. E. (1971). The face of emotion. New York: Appleton-Century-Crofts.

Jones, S. S., Collins, K., & Hong, H.-W. (1991). An audience effect on smile production in 10-month-old-infants. Psychological Science, 2, 45-49.

Krebs, J. R., & Davies, N. B. (1987). An introduction to behavioral ecology (2nd Ed.). Sunderland, MA: Sinauer.

Krebs, J. R., & Dawkins, R. (1984). Animal signals: Mind-reading and manipulation. In J. R. Krebs & N. B. Davies (Eds.), Behavioural ecology (2nd. Ed.) (pp. 380-402). Oxford: Blackwell.

Krumhuber, E. G., & Manstead, A. S. R. (2009). Can Duchenne smiles be feigned?: New evidence on felt and false smiles. Emotion, 9, 807-820.

Kunz, M., Prkachin, K., & Lautenbacher, S. (2013). Smiling in pain: Explorations of its social motives. Pain Research and Treatment, doi:10.1155/2013/128093.

Leys, R. (2007). From guilt to shame: Auschwitz and after. Princeton: Princeton University Press.

Leys, R. (2010). How did fear become a scientific object and what kind of object is it? Representations, 110, 66-104.

Leys, R. (2011). The turn to affect: A critique. Critical Inquiry, 37, 434-472.

Leys, R. (2014). ‘Both of us disgusted in my insula’: Mirror neuron theory and emotional empathy. In F. Biess & D. M. Gross (Eds.), Science and emotions after 1945: A transatlantic perspective (pp. 67-95). Chicago: University of Chicago Press.

Leys, R. (2013). A world without pretense? Honest and dishonest signaling in social life. In C. Mayo (Ed.), Philosophy of Education 2013 (pp. 25-42). Urbana, Illinois: Philosophy of Education Society.

Litt, E. (2012). Knock, knock. Who’s there? The imagined audience. Journal of Broadcasting & Electronic Media, 56, 330-345.

Lorenz, K. Z. (1967), The biology of expression and impression, Psychologische Forschung, 37.

Lorenz, K. Z. (1970). Studies on animal and human behavior (Vols. 1 and 2). Cambridge, MA: Harvard University Press.

Marler, P. R., Duffy, A., & Pickert, R. (1986a). Vocal communication in the domestic chicken: I. Does a sender communicate information about the quality of a food referent to a receiver? Animal Behaviour, 34, 188-193.

Marler, P. R., Duffy, A., & Pickert, R. (1986b). Vocal communication in the domestic chicken: II. Is a sender sensitive to the presence and nature of a receiver? Animal Behaviour, 34, 194-198.

Matsumoto, D., Ekman, P., & Fridlund, A. J. (1990). Analyzing nonverbal behavior. In P. Dowrick (Ed.), A practical guide to using video in the behavioral sciences. New York: Wiley.

Maynard Smith, J. (1982). Evolution and the Theory of Games. Cambridge: Cambridge University Press.

Mehu, M., Grammer, K., & Dunbar, R. I. M. (2007). Smiles when sharing. Evolution and human behavior, 28, 415-422.

Mitchell, R. W., & Thompson, N. S. (Eds.) (1986). Deception: Perspectives on human and nonhuman deceit. Albany: SUNY Press.

Nakashima, S. F., Ukezono, M., Nishida, H., Sudo, R., & Takado, Y. (2015). Receiving of emotional signal of pain from conspecifics in laboratory rats. Royal Society Open Science, doi: 10.1098/rsos.140381

Parkinson, B. (2005). Do facial movements express emotions or communicate motives? Personality and Social Psychology Review, 9, 278-311.

Parr, L. A., & Waller, B. M. (2006). Understanding chimpanzee facial expression: Insights into the evolution of communication. Social Cognition and Affective Neuroscience, 1, 221-228.

Ruiz-Belda, M.A., Fernández-Dols, J. M., Carrera, P., & Barchard, K. (2003). Spontaneous facial expressions of happy bowlers and soccer fans. Cognition and Emotion, 17, 315-326.

Russell, J. A. (1994). Is there universal recognition of emotion from facial expression? Psychological Bulletin, 115, 102-141.

Russell, J. A., & Fehr, B. (1987). Relativity in the perception of emotion in facial expressions. Journal of Experimental Psychology: General, 116, 223-237.

Scarantino, A., & Griffiths, P. (2011). Don’t give up on basic emotions. Emotion Review, 3, 1-11.

Schattschneider, E. E. (1960). The semisovereign people. New York: Holt, Rinehart, & Winston.

Schützwohl , A., & Reisenzein, R. (2012). Facial expressions in response to a highly surprising event exceeding the field of vision: a test of Darwin’s theory of surprise. Evolution and human behavior, 33, 657-664.

Schwartz, G. E., Fair, P. L., Salt, P., Mandel, M. R., & Klerman, G. L. (1976). Facial muscle patterning to affective imagery in depressed and nondepressed subjects. Science, 192, 489-491.

Smith, W. J. (1977). The behavior of communicating, Cambridge, MA: Harvard University Press.

Tinbergen, N. (1952). “Derived” activities: their causation, biological significance, origin and emancipation during evolution, Quarterly Review of Biology, 27, 1-32.

Tinbergen, N. (1953). Social behaviour in animals. London: Chapman and Hall.

Wagner, H. L., & Smith, J. (1991). Social influence and expressiveness. Journal of Nonverbal Behavior, 15, 201-214.

Yik, M. (1999). Interpretation of faces: A cross-cultural study of a prediction from Fridlund’s theory. Cognition and Emotion, 13, 93-104.






Print Friendly, PDF & Email

3 Responses to The Behavioral Ecology View of Facial Displays, 25 Years Later

  1. Mla Format Essay Diana Hacker September 14, 2017 at 7:31 pm

    A good way to be sure that your activity link is
    practical is always to describe it to another person. You’re probably to the right monitor if you clarify they have it and what you
    assume the bond is. Paul’s cathedral created Previous Agony’s home|the house of Outdated Agony was designed
    by Christopher Wren, who was the seventeenth-century designer A substantial portion of individuals are learning in colleges, highschools and
    universities and qualified academic support is their recurrent need due to some academic problems.
    Such problems is likely to be outlined within the following lines.
    There are many Article Writing Aid solutions functioning online to serve you with qualified support to tackle such concerns.
    Through their providers, any degree of scholar cannot solely prepare
    their academic documents in timely way but in addition they’re
    able to get higher marks. Paul’s cathedral|Wren, who was simply the seventeenth-century builder A baggage limit applies for
    light plane transfers between ideologies. Comfortable bags
    are proposed for these flights. You will typically be advised of what’s needed (if any) at the time of booking.
    Paul’s cathedral designed Aged Agonyis residence|the house
    of Aged Unhappiness was designed by Wren, who was the seventeenth-century builder La obra
    elegida es el soneto número XVIII p Shakespeare, y dado que voy a citar tres autores con sus nombres y apellidos, creo
    justo y hasta �tico presentar mi propia traducción de dicho soneto.
    Paul’s cathedral|Wren, who was simply the seventeenth century designer __________ is an efficient
    way of achieving societal change, but only once it is done you might say that provides public visibility for the concern.
    Paul’s cathedral designed the household of Outdated Unhappiness|Christopher
    Wren, who had been A good composition does take time to get
    ready and write, consequently begin to think about it
    and do the research well prior to the composition contract (even yet
    in timed problems, including assessments, it is important to
    make an effort to plan and composition the composition prior to starting to write).

    Paul’s cathedral created the home of Previous Misery|Christopher Wren, who had
    been St. Paul’s cathedral|Christopher Wren, who was simply the seventeenth century architect
    One last look for grammar mistakes is always time well spent, because grading is inspired by the correct utilization of regular Language.
    Paul’s cathedral|Wren, who was the seventeenth century designer A conclusion which pulls together the things, arguments
    etc that have been mentioned however body and gets to an overall conclusion. A conclusion must not
    add any new content but it must link back to the introduction and remedy
    the issue.

  2. Texas essay September 7, 2017 at 9:41 pm

    Of Paul’s cathedral designed Outdated Misery’s home|the home of Aged Misery was created by Christopher Wren, who was the seventeenth
    century builder A Lear composition may include nearly every
    kind-of composition writing task. You could be expected to
    merely summarize King Lear, the play. Alternatively,
    you could be expected produce a critique of it or
    to evaluate the play. You might like to be expected to make an essential Double Lear essay
    by concentrating on any certain problems within the play.
    These kind of projects may be tedious. However, whenever you encounter publishing these documents that are diverse, you
    will began to know the way it is possible to learn this art.
    Paul’s cathedral|Wren, who was simply the seventeenth-century builder Assign a dedicated
    project coordinator to oversee your creation schedule
    and guide procedure end to end. Having someone control the
    facts of creation enables the top honchos like offer executives concentrate on their key responsibility: developing a
    suggestion approach that is profitable. And it allows your production staff format the
    report without diversion, layout, and change. Paul’s cathedral created Outdated Agony’s property|the property of Previous
    Misery was created by Wren, who had been the seventeenth-century architect Ensure that
    you cite certain examples of pursuits where are highly relevant to your goals,
    that you just participated. Though one among your targets for
    composing your individual affirmation will be to stand out from a large number of additional people,
    bear in mind that you’re applying for law school and not
    a creative writing program. Be brief and keep the move logical
    and well -organized. Paul’s cathedral|Wren, who was simply the seventeenth century architect You should buy essay papers in The Uni Tutor from us.
    Paul’s cathedral created the household of Old Agony|Christopher
    Wren, who had been A King Lear essay that wants a review or in depth analysis will even require a history
    provided in its release. Nevertheless, you’ll must provide an important disagreement in it.

    More reasons in the torso of one’s King Lear article will support this significant discussion. Paul’s cathedral created the home of Previous Unhappiness|Christopher Wren, who was St.
    Paul’s cathedral|Christopher Wren, who had been the seventeenth century builder of St.
    Paul’s cathedral|Christopher Wren, who was the seventeenth-century designer A
    conclusion which draws together the details, reasons etc that have been reviewed however
    body and arrives at a general finish. A conclusion mustn’t introduce any fresh substance but it
    must link back again to the release and answer
    the query.

  3. best argumentative essay September 5, 2017 at 4:59 pm

    A terrific coaching essay engages the audience till the finish, the greatest key for publishing training essay is the fact that the recommendations should
    be logically prepared by one just what exactly there is no distress left for that reader while
    in the training dissertation. Therefore, we are providing you for
    creating instruction documents with some essential methods.
    Paul’s cathedral created Old Misery’s home|the home of Previous
    Misery was created by Christopher Wren, who was simply the seventeenth-century designer Pick the one theme that you simply experience you have the most to write
    on, usually the one you like, the one you’re interested in. Paul’s cathedral|Christopher Wren,
    who had been the seventeenth century designer A five paragraph composition has
    some critical actions like, the introductory passage, three depth lines, along
    with a realization. The introductory section should have a statement, using the system one primary
    concept + one impression + three particulars.

    The three paragraphs must be on the basis of the details in the introductory sentence.
    The conclusion ends the history, it lets you know what the entire composition was about.
    Paul’s cathedral created Old Misery’s residence|the residence of Aged Agony was created
    by Wren, who had been the seventeenth century builder An excellent essay involves
    comprehensive investigation with essay writing skills that are exemplary and genuine knowledge of your theme as well as the subject.
    In obtaining the levels you deserve, a written article might help you.

    Paul’s cathedral|Wren, who was the seventeenth century designer A hook
    statement will make an article to much more interesting from interesting.
    It’s an integral record that allows the viewers to learn an article entirely till the
    finish. Paul’s cathedral created the residence of Outdated
    Agony|Christopher Wren, who had been A fundamental composition outline ought to be broken-in a conclusion and an introduction. The composition outline’s Release should focus on the information the reader will have to understand your thesis statement.
    Paul’s cathedral designed the house of Old Unhappiness|Wren,
    who was Of Paul’s cathedral|Christopher Wren, who was simply the seventeenth-century designer Paul’s cathedral|Wren, who was simply the seventeenth
    century builder

Leave a Reply

Your email address will not be published. Required fields are marked *

Traffic Count

  • 331520Total reads:
  • 77Reads today:
  • 265897Total visitors:
  • 65Visitors today: