The evolutionary bases of social and moral emotions:Dominance, submission, and true love

Ross BuckUniversity of Connecticut

Storrs, CT USA

Keywords

Acknowledgements

Paper to be presented at the9th Sydney Symposium of Social Psychology

The evolution of the social mind: Evolutionary psychology and social cognition

14-16th March 2006

In the first edition of Human Motivation and Emotion (1976; 1988a), this author pondered the question of whether human nature is good or evil: whether in the “state of nature” people lived together peacefully without leaders as imagined by Rousseau and

John Locke; or whether they were aggressively engaged in a constant “war of all against all” as argued by Thomas Hobbes. My answer, after considering studies of exploratory

behavior and conformity/obedience, was that the central facts of human nature are in fact powerful drives of curiosity directed toward the achievement of competence, and a great

susceptibility to social influence: “human beings are passionate creatures who are powerfully oriented toward distant goals and strongly influenced by their fellows. From these qualities come much that is good in humanity, and much that is evil” (Buck, 1976, p. 413). Thirty years on, it is perhaps appropriate to revisit this conclusion in light of the

enormous advances made in the interim in the literatures in evolutionary theory and affective neuroscience. This paper considers the deep evolutionary roots of motivation,

emotion, and communication; with the goal of examining the essential natures of cooperation and competition from the viewpoint of developmental-interactionist (DI)

theory (Buck, 1985; 1999). I first consider the definition and evolution of communication, with particular attention to the “selfish gene” view and its apparent

implication that genuine prosociality—“true love”—is impossible at the biological level. I counter that an alternative “communicative gene” view implies that a biologically-based

capacity for genuine prosociality is not only possible but indeed is required to explain basic facts of the role of communication in evolution. I then summarize recent evidence that prosocial motivational-emotional systems are in fact clearly present in the structures

and chemicals of the brain, and suggest that in human beings powerful attachment systems underlie systems of higher-level social and moral emotions that emerge naturally

and effortlessly from interaction over the course of development.

Genes, Communication, and Prosociality

The Evolution of Communication. Symbolic, spontaneous, and pseudospontaneous communication. “Communication”

is here defined following E. O. Wilson (1975) as occurring “whenever the behavior of one individual (the sender) influences the behavior of another (the receiver)…behavior can be defined as communicative to the extent that it reduces uncertainty in the behavior of another” (Buck, 1984, p. 4). In DI theory, human communication proceeds in two simultaneous “streams:” one voluntary and symbolic, the other automatic and spontaneous (Buck, 1988b, 1989, 1994; Buck & VanLear, 2002). Voluntary symbolic communication is learned and culturally patterned, its elements are symbols which bear an arbitrary relationship to the referent (for example, the word “tree” in English is arbitrarily related to the object it represents), and its content consists of falsifiable statements or propositions. A sender voluntarily encodes information into symbols, which are transmitted to a receiver who, assuming the receiver has learned the symbols and their rules for combination (vocabulary and grammar), can decode the message and thereby come to know the information. Both sender and receiver must have learned a similar language in order for symbolic communication to be successful.

In contrast, spontaneous communication is biologically structured in both its sending and receiving aspects. An internal motivational/emotional state of the sender is

automatically and effortlessly expressed in an evolved display, which given attention is picked up by the receiver and “known” directly via evolved preattunement as a motivational/emotional response in the receiver. The display is not a symbol, but rather is a sign of the internal motivational/emotional state. A sign bears a natural relationship to the referent—it is an externally accessible aspect of the referent, as in smoke being a sign of fire—so that if the sign is present the referent must be present by definition. Therefore symbolic communication is not propositional in that by definition it cannot be false.

These two simultaneous streams of communication coexist in virtually every human communicative exchange, although their relative importance will vary. In formal and structured situations, such as a lecture, symbolic communication tends to dominate but spontaneous communication plays a subsidiary albeit important role in conveying for example the charisma of the speaker and enthusiasm of the audience. This domination of symbolic communication is often aided by the architecture of the lecture room, with the speaker elevated and facing the audience, and the audience oriented toward the speaker. In a less structured seminar, the role of spontaneous communication is enhanced, often in part by a face-to-face orientation of the seminar participants. In the “throes of passion,” spontaneous communication comes to predominate: symbolic and rational considerations are set aside in favor of strong emotions: love, ecstasy, rage, misery, terror.

The symbolic-spontaneous mix also varies with the intimacy of personal relationship of sender and receiver. All else equal, symbolic communication predominates in formal relationships where the actors do not know one another well, but as personal relationships develop and become more intimate, the relative importance of spontaneous communication tends to increase. The symbolic-spontaneous mix also varies over the course of human development: the newborn is primarily a spontaneous communicator, but as an infant grows into a toddler and comes to learn language, symbolic communication becomes more and more important (Buck, 1982). Finally, the symbolic-spontaneous mix varies along the evolutionary scale: as species increase in complexity, relatively inflexible spontaneous communication systems interact progressively more extensively with general-purpose symbolic communication systems, so that communication becomes progressively more flexible. This progressive evolution of increased behavioral plasticity has been termed anagenesis (Gottleib, 1984).

There is evidence that the individual has considerable “voluntary” control over the display, in that it is possible for a sender to show motivational and emotional displays that are not really present as internal states. Arthur VanLear and I termed this “pseudospontaneous communication,” because while it is voluntary from the sender’s point of view, it uses the display mechanism that can stimulate preattunements in the receiver, so if the receiver is taken in it is as if it were a veridical display. A charismatic sender, for example, can successfully “push the buttons” of an audience, and persuade by manipulating others’ emotions (Buck & Van Lear, 2002). This voluntary expression of a display was termed “voluntary expression formation” in analyses of brain mechanisms of primate audiovocal communication by Jurgens (1979) and Ploog (1981). The question of veridical versus manipulative displays is a fundamental issue in the evolution of communication, as we shall see.

Classical ethology: communication and group selection. The classical analysis of the evolution of communication stems from Charles Darwin’s theory of evolution, and particularly Expression of the Emotions in Man and Animals (1872/1998). Darwin

suggested that emotional displays can have adaptive value in social animals because they reveal inner states of the responder that are useful for social co-ordination, including for example aggressive dominance, fearful submissiveness, and sexual readiness. This implies that the inner state of the responder (sender) must be associated with external expression, and that the receiver must be able to “pick up” the expression via sensory cues: postures, facial expressions, pheromones. Darwin's thesis requires that sending and receiving mechanisms co-evolve—evolve simultaneously with one another—for the adaptive value of a system of communication to be realized. This analysis reflects the features of spontaneous communication as defined previously.

Ethologists including Lorenz and Tinbergen agreed that the communication of certain motivational-emotional states is adaptive (Hauser, 1986). Those individuals who show evidence of that state in behavior will tend to be favored. Over the generations these behaviors can become “ritualized” into expressive displays. Similar reasoning was applied to the evolution of receiving mechanisms: individuals who respond appropriately to these displays would tend to be favored, so that the perceptual systems of species members can become “preattuned” to the pickup of these displays. In this way, displays and preattunements coevolve as aspects of a system of spontaneous communication.

Some of Darwin’s successors advanced the notion that behaviors, including communicative behaviors, evolve as group-level adaptations (Kreft, 2004a). It seemed intuitively logical that successful groups would foster qualities in group members that would support and sustain that success. For example, Dobzhansky (1937) suggested that species maintain genetic diversity to cope with environmental change; Allee (1951) held that dominance hierarchies function to minimize within-group conflict; and Wynne-Edwards (1962) maintained that individuals restrain from breeding to avoid population die-offs.

The “selfish-gene” critique of group selection. In the 1960’s, such group-level thinking was criticized as naïve, illogical, and unscientific. G. C. Williams (1966) argued that the selection of individual-level benefits is more immediate and effective than the selection of group-level benefits, and that group-level selection should only be invoked if individual-level benefits have been ruled out (Kreft, 2004a; Redfield, 2002). The classical view was further challenged when selection was interpreted as operating, not at the level of the individual or group, but rather at the level of the gene (Hauser, 1996). In The Extended Phenotype (1982), Richard Dawkins derided what he termed “sloppily unconscious group-selectionism” (p. 6). He argued that the ultimate unit of evolutionary selection is the active, germ-line replicator. A replicator is “anything in the universe of which copies are made,” an active replicator is “any replicator whose nature has some influence over its probability of being copied,” and a germ-line replicator is a “replicator that is potentially the ancestor of an indefinitely long succession of descendent replicators” (1982, p. 83). Dawkins argued that the only active replicator lasting across evolutionary timescales is the gene, so that the “selfish gene” is the unit of selection, a replicator motivated only to make copies of itself. Fitness was seen as based upon the survival, not of the individual organism or the group, but rather upon inclusive fitness: the survival of the genes.

Dawkins noted that, for some purposes, it is convenient to think of the effects of replicators as being packaged together in discrete “vehicles” such as individual organisms or groups, but such vehicles are not units of selection because their influence does not

extend over evolutionary timescales: “to the extent that active germ-line replicators benefit from the survival of bodies in which they sit, we may expect to see adaptations that can be interpreted as for bodily survival…To the extent that active germ-line replicators benefit from the survival of bodies other than those in which they sit, we may expect to see ‘altruism,’ parental care, etc. …But all of these adaptations will exist, fundamentally, through differential replicator survival” (Dawkins, 1982, pp. 84-85. Italics in the original). All in all, the critique of the naïve appeal of group-level selection was devastating, and effectively “outlawed group-level thinking” in evolutionary biology (Kreft, 2004a, p. 267).

Selfishness and communication. The “selfish gene” critique extended to the understanding of communication as well, and specifically to the idea that accurate communication is adaptive. Instead, it was argued that selection would actually operate against those who show veridical displays that are predictive of their true inner states and probable behaviors, and suggested that communication is actually a means by which one animal exploits another: “a means by which one animal (the 'actor') exploits another animal's (the 'reactor's') muscle power” (Krebs and Dawkins, 1984, pp. 380-381). Krebs and Dawkins argued that displays evolve via selection for effective manipulation, defined as the sender “actively changing the victim's (sic) behavior” (1984, p. 383).1 In this regard, Dawkins and Krebs (1978) suggested an analogy between animal communication and media advertising, where the object is persuasion rather than transfer of information. This manipulative communication corresponds to pseudospontaneous communication as defined previously, where the display does not in fact truly reflect an internal state but rather is “put on” voluntarily by the sender, but it can activate the preattunements in the receiver and therefore be emotionally compelling.

Krebs and Dawkins (1984) suggested that mind-reading on the part of the receiver--interpreting the actual internal state and predicting the behaviors of other animals--is the counterpart to manipulation on the part of the sender. The sender's counter-response to mind-reading may involve concealment (a poker-face) and active deception (simulating, qualifying or falsifying one's display). The authors presented manipulation and mind-reading as “intimately locked together in evolutionary arms races and feedback loops…Mind-reading and manipulation coevolve, and signals are the result of this coevolution” (Krebs and Dawkins, 1984, p. 389).

Multilevel selection theory. Sober and D. S. Wilson (1998; D. S. Wilson & Sober, 1994) suggested an alternative to the selfish gene position based upon the definition of the unit of evolutionary selection. They suggested that evolution can take place at multiple levels, including the gene, the individual organism, and the group. However, this view was criticized by Dawkins (1994) because individuals and groups do not exist across evolutionary timescales. He argued that individuals and groups are vehicles of selection that are "not something fundamental" in evolution (Dawkins, 1994, p. 617), and that although group selection is possible theoretically, it is not important in nature.

Altruism.

1 I use the terms "sender" and "receiver" in place of Krebs and Dawkins' (1984) "actor" and "reactor."

Kin selection and reciprocal altruism. Altruism is defined as the sacrifice of one’s own genetic fitness in favor of benefiting the genetic fitness of another. The phenomenon of altruism poses a fundamental problem for evolutionary theory. Dawkins (1989) argued that the “law of ruthless selfishness” governing the selection of genes implies that true altruism is impossible: “Be warned that if you wish, as I do, to build a society in which individuals cooperate generously and unselfishly towards a common good, you can expect little help from biological nature. I am not advocating morality based on evolution. Let us try to teach generosity and altruism, because we are born selfish (p. 3).”

Despite this, there are apparent examples of unselfish, cooperative, and even altruistic behavior that must be taken into account. It is widely agreed that, under conditions of kinship and/or reciprocity, mutually cooperative communication can foster the inclusive fitness of the altruist, and therefore it can be favored by selection. In kin selection, one helps genetically-related others, therefore indirectly preserving the genes shared with the relative (Hamilton, 1963; 1964). In reciprocal altruism, one helps with expectation of return (Trivers, 1971). Because such behavior fosters the survival of the altruist’s own genes via inclusive fitness, such apparent altruism is actually selfish, and as such it does not contradict the selfish gene hypothesis (Krebs and Dawkins, 1984). Evidence that human beings sometimes appear to engage in altruistic behaviors despite considerations of kinship and reciprocity being apparently absent has been answered by the argument that humans are special because, as Dawkins noted, generosity and altruism can be learned. For example, Rachlin (2002XX) argued that apparent altruism can be a consequence of a learned commitment to an altruistic pattern of behavior.2 Such learning must necessarily go against the naturally selfish tendencies of “biological nature.”

Quorum-sensing in bacteria. Recently, evidence relevant to the evolution of altruism has come from an unexpected source: microbiology. In Animal Aggregations (1931), W. C. Allee noted self-organizing activity in simple creatures, including bacteria. Bacteria are prokaryotes—essentially bags of DNA—that lack the nuclei, mitochondria, and organelles associated with the eukaryotic cells that make up all complex multicelled life. Despite their relative simplicity, recent studies have discovered surprising complexity and sophistication in the behavior of bacteria, including intra- and intercellular communication resulting in behavior suggestive of intelligence and memory (Hellingwerf, 2005), cooperation and altruism (Griffin, West, & Buckling, 2004; Kreft, 2004a, 2004b), and even “social intelligence” (Ben-Jacob, Becker, Shapira, and Levine, 2004). In addition, there is evidence that prokaryotes lived socially from the beginning.The most ancient known organisms on Earth are fossilized colonies of cyanobacteria termed stromatolites: stromatolites were abundant 3.5 billion years ago and exist today living in suitable environments in Australia and Baja California (Olson, 1989). Today, cyanobacteria are responsible for the pond scum that forms on stagnant water. In stromatolites, these ancient, relatively uncomplicated creatures self-organize and self-

2 It is noteworthy that in his Behavioral and Brain Sciences paper "Altruism and selfishness," Howard Rachlin used the word "choice" 80 times, while "emotion" did not appear. In contrast, a paper in the same journal, "Empathy: Its ultimate and proximate bases" by Stephanie D. Preston and Franz B. M. de Waal (2001), used "emotion" 139 times and "choice" once. Clearly, these articles represent fundamentally different ways of looking at the phenomena of empathy and altruism, and each does not address the positions of the other (Buck, 2002).

configure to create a working community that recovers from damage, using systems of communication that are not well understood.

Recent studies have elucidated mechanisms by which such bacterial social behavior is coordinated. Many if not most species of bacteria exhibit quorum-sensing: mechanisms for recruiting the mass production of molecules or engaging in other collective activities beneficial to the bacteria. The bacteria are quiescent until a critical mass of individuals—a “quorum” of millions or billions—has assembled, and they then produce the molecule en mass in a useful concentration (Swift et al., 1996; Waters & Bassler, 2005).

It was thought until recently that quorum sensing was restricted to relatively few species of bacteria, and served restricted functions. One well-described example is the marine bioluminescent bacterium Vibrio fischeri. This bacterium lives freely in a planktonic state, and also exists in symbiotic relationship with certain fish and squid (i.e., the Hawaiian squid Euprymna scolopes), causing luminescence that functions to attract food and camouflage the squid in moonlight (Waters & Bassler, 2005). In the laboratory, a growing colony of V. fischeri remains dark until a relatively high density of individuals is achieved, at which point luminescence increases rapidly. This phenomenon is caused by the action of signal molecules, which increase in concentration with an increasing number of individuals. The signal molecule responsible for the activation of luminescence in V. fischeri was identified in 1981, and the genetic system analyzed in 1983 (Eberhard et al., 1981; Engebrecht, Nealson, & Silverman, 1983). When the level of signal molecules reaches a threshold, they enable proteins called LuxR to bind to specific genes within the individual cells. A molecular apparatus “turned on” by the interaction of LuxR and genes generates the light simultaneously in many individual bacteria (Fuqua, Winans, & Greenberg, 1996; Greenberg, 1997; Schaefer et al., 1996; Waters & Bassler, 2005).

One of the general problems of a system that works by cooperation is the presence of cheaters who attempt to cash in on the benefits of the system without doing their part (Travisano & Velicer, 2004). In the symbiotic relationship between V. fischeri and E. scolopes, the bacteria receive rich nutrients that allow proliferation in exchange for the benefit of giving light at the appropriate time to camouflage the squid. There are individual bacteria, however, that potentially could bask in the benefits of the hospitality of E. scolopes without incurring the metabolic costs of actually lighting up. If that strategy were to succeed, the lights would soon go out. As it is, V. fischeri mutants who are incapable of luminescence are outcompeted by their luminescent cousins in the squid host: E. scolopes appears to possess a “policing” mechanism that distinguishes cells incapable of luminescence and acts to eliminate cheaters (Waters & Bassler, 2005; Visick, Foster, Doino, McFall-Nagai, & Ruby, 2000).

Interest in quorum-sensing within microbiology has exploded in the past decade: “the number of known regulatory systems and the diversity of phenomena regulated are growing dramatically, and it now appears that most bacteria possess at least one quorum-sensing system” (Redfield, 2002, p. 365). While the functions of these systems are extraordinarily diverse, the systems by which quorum-sensing is based are surprisingly uniform. Quorum-sensing works by a bacterium’s release of an autoinducer molecule into the environment. These constitute signals or displays, and typically involve amino acids or peptides functioning as pheromones (Gallio, Sturgill, Rather, & Kylsten, 2002). The bacterium also has the capacity to sense the concentration of this autoinducer in the

environment. If the concentration exceeds a threshold, the expression of genes within the bacterium is altered, producing a variety of effects: motility, swarming, pigment formation, etc. (Redfield, 2002). For example, quorum sensing is involved in some bacterial infections, such that when a quorum is attained, the regulation of bacterial genes is changed so that toxic “virulence factors” are produced. In cystic fibrosis, bacteria produce a “biofilm” when they reach a certain concentration, a tough shell that protects the bacteria from attack from the immune system of the victim and/or antibiotics. The bacteria then can reproduce, produce toxins, and damage tissues in relative safety (Riedel & Ebert, 2002). Biofilms have been characterized as “microbial societies with their own defense and communication systems” (Costerton, Stewart, & Greenberg, 1999, p. 1322). One feature of quorum sensing is that the bacteria may be quiescent—not activating a full immune system response—until their number is sufficient to overwhelm the victim.3

Altruism and quorum-sensing. The production by a bacterium of an autoinducer, and its responsiveness to the environmental concentration of the autoinducer, has all of the qualities of spontaneous communication as defined previously. Both the display (autoinducer production) and preattunement (responsiveness) are biologically-based, the autoinducer is a sign of the referent (concentration of individuals), and the communication is in no way intentional and is nonpropositional. The question whether quorum-sensing in fact represents a system of communication to promote collective action useful to the group is relevant to the question of cooperation and altruism in evolutionary biology. In everyday language, altruism is typically related to a concern for the welfare of others and the common good. In the context of evolutionary theory, these are not considered: “Evolutionary altruism…does not require memory of past

interactions, recognition of individuals, sophisticated interactions or behavioral repertoires, or direct interactions between individuals. It is therefore the simplest form of altruism” (Kreft, 2004b, p. 2751).

In quorum-sensing, the concentration of autoinducer ordinarily exceeds threshold only when the population of cells reaches a certain density, and the most widespread interpretation of the quorum-sensing phenomenon is that it functions to sense population density. Redfield (2002) has suggested that this interpretation of quorum-sensing is incorrect for reasons that echo the selfish gene critique of group-level selection: “the difficulty of maintaining genes for cooperative strategies in genetically mixed populations makes this a notoriously weak and controversial mode of selection… (p. 365).” Furthermore, “those explanations that mesh well with our preconceptions are often uncritically accepted while those that do not are scorned…perhaps because we are social animals, we find the idea that bacteria have evolved communication and cooperation very appealing” (Redfield, 2002, p. 369). Redfield suggests that quorum-sensing is actually a side effect of diffusion sensing: the ability to detect whether secreted molecules move away from the cell.

3 The mechanism of quorum sensing is of great interest. If therapeutic agents can disrupt bacterial communication, the virulence of the bacteria can be greatly reduced (Costerton, et al., 1999; Foster, 2005). Moreover, because such treatments would target the mechanism of communication—in effect disrupting the dyad-level communicative relationship between bacteria rather than the bacteria as individuals—it would be less likely to leave drug-resistant individuals behind to reproduce and eventually create strains of drug-resistant bacteria (Buck & Ginsburg, 1991).

Despite these objections, there is concrete evidence of individual sacrifice for group benefit in microorganisms (Walters and Bassler, 2005). One example involves the slime mold Dictyostelium discoideum. At one stage in its life cycle, slime molds exist as unicellular amoebae feeding on bacteria. At this stage the amoebae exhibit positive chemotaxis (attraction) vis a vis their prey and negative chemotaxis (repulsion) vis a vis one another.4 The attractant is folic acid, an essential vitamin that the amoebae obtain from ingesting bacteria. At the same time, the amoebae release an unidentified substance that keeps other amoebae of the same species away: functioning to "aid dispersion of the amoebae and therefore permit better utilization of the environment" (Lackie, 1986, p. 234). Thus this simple organism demonstrates a sort of "threat display" that may fulfill functions analogous to those ascribed to territorial displays in more complex creatures (Eibl-Eibesfeldt, 1975).

As bacteria in the environment are consumed, the amoebae begin to starve, the negative chemotaxis to other amoebae ceases, and a positive chemotaxis begins. Within 4-6 hours, the individual amoebae begin to move toward aggregation centers, which apparently contain those individuals whose positive chemotaxic systems were first "turned on" by starvation. The chemical attractant responsible for aggregation is cyclic adenosine monophosphate (cAMP), to which individual amoebae respond by themselves releasing cAMP, thereby relaying the signal (Lackie, 1986). The initial phase involves individual movement toward the center based upon perception of the dynamic gradient of cAMP (interestingly, the amoebae do not respond to a static gradient of cAMP: Vicker, Schill & Drescher, 1984). This phase is followed by a phase of streaming based upon "nose to tail" contact following, facilitated by specialized contact sites which appear as adhesion molecules now defining the front and rear of the individual amoebae (Gerisch et al, 1975; 1980). It is noteworthy that competition between aggregation centers occurs, based in part upon gradient interactions and in part apparently upon "dominance" by older, more established, aggregation centers; although the mechanism of the latter is unknown. It is also significant that, if mixed experimentally, individual amoebae of different species aggregate independently (Lackie, 1986).

The aggregation center next forms a multi-celled slug, or grex, in which the cells are derived from individual amoebae. The grex moves in a looping motion like an inch-worm caterpillar. The size of the grex in D. discoideum ranges from 100,000 to 200,000 individual cells, so it must solve significant problems of coordination: as Lackie put it "100,000 dissenting voices are scarcely a recipe for united action" (1986, p. 239). In fact, the grex demonstrates coordinated responses to light, temperature, humidity, and oxygen. The mechanism of coordination is not well understood, although gradients of cAMP within the grex are involved, and lateral adhesions between individuals in addition to the nose-to-tail joining may confer some mechanical coordination.

The grex moves from the domain of the individual amoebae in damp forest litter, which is inappropriate for dispersal, up surface layers toward the light in a journey that may take many days. The sensory analysis of the environment presumably takes place in the front tip of the grex, and it is the cells in this area that become anchored and "altruistically" die to form the cellulose stalk of a fruiting body (Strassmann, Zhu, & Queller, 2000). The stalk formation is altruistic on the part of these individuals in the

4 Chemotaxis is "the directed movement of a cell (or organism) in response to a chemical substance in the environment" (Lackie, 1986, p. 220).

technical evolutionary sense: they "surrender...personal genetic fitness for the enhancement of personal genetic fitness in others" (E. O. Wilson, 1975, p. 106), or behave "to increase another such entity's welfare at the expense of its own" (Dawkins, 1976, p. 4). Cells at the rear of the grex form a mass that climbs to the top of the stalk to become individual spores, which are released into the environment. Given favorable conditions, they germinate into individual amoebae, and begin the life cycle again.

In D. discoideum, the development of fruiting bodies is initiated by cAMP and Differentiation-Inducing Factor (Town, Gross, & Kay, 1976; Town and Stanford, 1979). In the soil-dwelling bacterium Myxococcus xanthus, a similar process involves quorum sensing (Clarke, 1981; Losick & Kaiser, 1997; Shimkets, 1999). In both cases, “spore development requires a large percentage of the population to undergo a lethal differentiation event that leads to structures whose function is to promote spore generation and dispersion” (Waters & Bassler, 2005, p. 336). These indeed appear to be examples of altruism—in the technical evolutionary sense—at the microbial level: how can this be reconciled with the powerful and compelling gene-selectionist account of evolution?

The Communicative Gene Hypothesis. Communication and the unit of selection. The issue of the possibility of "true"

altruism turns on the issue of the unit of selection: whether the unit of selection is the individual gene as "selfish gene" proponents propose, or whether evolution can involve the selection of units above the level of the individual gene. A possible solution is that communicative relationships between genes can be replicators; units of selection in evolution (Buck, 2002; Buck & Ginsburg, 1991; 1997a; 2000).

It is possible to retain a gene-selectionist position without a largely unexamined accompanying assumption of genetic atomism: genes are selected in isolation from other genes: "selection purely at the level of the individual gene" (Dawkins, 1989, pp. 84-85). Buck & Ginsburg (1991) noted that genes do not function alone. Rather, genes function in company with other genes, and more specifically, genes function by communicating with other genes. This is not a controversial contention, but its implications regarding the unit of selection have perhaps not been fully appreciated.

The genes that underlie a trait constitute the genotype, while the phenotype is the expression of that trait in the qualities and behaviors of the individual organism that are actually selected in the course of evolution. Genes are selected by proxy, via the selection of the phenotype. In the case of communication, the phenotype subject to selection is functional communication accuracy, the accuracy of gene A sending and gene B receiving (Ca b). If individuals who communicate effectively have a greater chance of survival, the sending accuracy of gene A to gene B and the receiving ability of gene B from gene A will be selected simultaneously. So, genes associated with a sending mechanism (for example controlling a bacterium’s release of an autoinducer molecule) and a receiving mechanism (controlling the capacity to sense the concentration of this autoinducer) must co-evolve. This is not “selection at the level of the individual gene,” but rather it is selection at the level of the communicative relationship of genes A and B, and the result is the display-preattunement relationship characteristic of spontaneous communication.

The elements of communication. A critical postulate of the communicative gene view is that in any system of interacting elements, communication involves both individual elements and the unique relationship between those elements relative to other elements. Thus, Ca b accuracy depends in part on the individual abilities of gene A to send (Sa) and gene B to receive (Rb), but it also depends in part upon the unique ability of gene A to send to gene B relative to other genes (Ua b). Similarly, the reciprocal ability of gene B to send to gene A depends in part on the ability of gene B to send (Sb) and gene A to receive (Ra), and in part to the unique ability of gene B to send to gene A relative to other genes (Ub a).

This breakdown of communication into sending effects, receiving effects, and unique relationship effects is a feature of the Social relations Model (SRM: Warner, Kenny, & Stoto, 19XX; Kenny, 1994). This defines the following as making up communication between genes A and B:

1. Ca b = (Sa + Rb + [Ua b] + error)2. Cb a = (Sb + Ra + [Ub a] + error)

These terms can be measured or estimated via round-robin designs in which communication between A and B are assessed in company with their communication vis a vis other elements (e.g., C, D, E, etc.). The SRM allows the analysis of the proportion of variance in total communication accuracy due to individual sending and receiving accuracies, and that due to the unique relationship between the individual elements plus error, which can be estimated in designs incorporating repeated measures.

To recapitulate, communication accuracy (Ca b) is the phenotype, the trait selected by evolution. The SRM shows that Ca b reflects not only the individual sending accuracy of A and receiving ability of B, but the unique relationship of A and B as well. This unique relationship (Ua b) is an aspect of the genotype which is selected by proxy via the selection of the phenotype Ca b. Studies using the SRM have consistently found that a substantial portion of variance if communication accuracy—up to 50% or more—is due to the unique relationship between individuals as opposed to individual sending and receiving abilities (Sabatelli, Buck & Kenny, 1986, Kenny, 1994).

Although the SRM was developed in the context of social psychology, the point regarding communication—that both individual and dyad-level components make up observed communication accuracy—is general, applying to any system of communicating elements including genes. It is possible in principle to measure the interaction between specific genes using the equivalent of a round-robin design: that is, controlling for their interactions with other genes, and therefore assessing unique relationship effects.

Implications for altruism: Relationships as replicators. This analysis suggests that the genotype (the replicator) includes potentially measurable relationship factors; that communicative relationships can be replicators in Dawkins’s (1982) sense. Communicative relationships arguably meet the criteria for being active germ-line replicators. A replicator is anything of which copies are made; an active replicator influences the probability of being copied; a germ-line replicator is an ancestor of descendant replicators; and replicators exist across evolutionary time scales. Communicative relationships (Ua b) and (Ub a) can be copied via the selection of the phenotype Ca b and Cb a; the nature of (Ua b) and (Ub a) can influence the probability of being copied; and these relationships can exist across evolutionary

timescales. Therefore, communicative relationships between genes can be active, germ-line replicators. Moreover, communicating genes are not necessarily within the same cell or organism. The quorum-seeking example demonstrates that genes in different individual bacteria can communicate via autoinducer molecules functioning as signs of population density: it involves displays in the sender and preattunements in the receiver.

There are many examples of specific communicative relationships that have existed across evolutionary timescales: indeed the ritualized displays associated the classical ethological view meet this criterion. Specific displays associated with dominance, submission, warning, courting, and bonding have existed across evolutionary timescales and in many species: for example, Livingstone, Harris-Warrick, and Kravitz (1980) demonstrated that serotonin injections in crayfish and lobsters produce characteristic dominance postures, while injections of octopamine (the phenol analogue of norepinephrine) produce subordinate postures, and the mating pheromone in the single-celled yeast Saccharomyces cerevisiae is a peptide molecule that closely resembles GnRH involved in mating in mammals, including human beings (Loumaye, Thorner, & Catt, 1982). Both of these imply a conservation of sending and receiving mechanisms functioning across an enormous span of time.

As noted, there is always the potential for cheaters to exploit a cooperative relationship, and the active mechanisms to respond to cheaters even in microbial life noted previously demonstrates that cooperation is never a simple matter. But, although manipulative communication clearly can be of selfish benefit, it does not follow that all communication is manipulative: veridical communication must also exist, and indeed, the presence of lying itself implies an underlying truth. Winston Churchill said in 1944: “truth is so precious that she should always be attended by a bodyguard of lies” (REFXX). This statement expresses a fundamental truth about truth and lying, but if there were not in fact an underlying truth the lie would not be necessary: a bodyguard of lies is pointless without a body to guard, as it were.

Summary and ImplicationsIn summary, the communicative gene hypothesis makes allowance for biologically-

based cooperative tendencies as well as selfish tendencies, suggesting that built-in tendencies for genuine altruism, self-sacrifice, attachment, and “true love” can be products of evolution alongside selfishness and calculations of self-interest. Indeed, when the nature of communication is taken into account, a gene-selectionist viewpoint arguably not only allows, but requires, true altruism. This conclusion is supported when we examine the motivational-emotional systems actually present in the brain. Largely separate from the debate about the selfish gene position and its implications for altruism, evidence has been accumulating in the neurosciences for the existence of powerful prosocial biological emotions (Buck, 1999, 2002). The existence of these prosocial neurochemical systems seems contrary to the notion that altruism is impossible at the biological level, and must be learned.

It is noteworthy in this regard that Darwin himself did not deny the possibility of biologically-based prosocial motives and emotions. Indeed, he argued that compassion and moral sensitivity—”social instincts and sympathies”—are critical to the social order. He wrote, “the emotion of love…is one of the strongest of which the mind is capable” (1872/1998, p. 212).

Prosocial Biological Emotions

Oxytocin and VasopressinMost emotions examined by contemporary emotion theories involve selfish and

individualistic concerns: the “primary affects” of Tomkins, Izard, and Ekman and Friesen (e.g., happiness, sadness, fear, anger, surprise, disgust) all promote the adaptation of the individual. However, approaches that analyze brain mechanisms of emotion have found powerful evidence of neurochemical systems that appear to underlie prosocial emotions (Buck, 1999; 2002; Buck & Renfro, 2006). Based upon early studies of the effects of brain stimulation, Paul D. MacLean, Detlev Ploog, and others argued that while some emotions are inherently selfish, directed toward the survival of the individual organism; others are inherently social, directed at the survival of the species (MacLean, 1993; Ploog, 1981).

More recently, specific central neurochemical systems have been identified that are strongly associated with prosocial attachment: social recognition, mating, parental nurturance, bonding, and play. These systems involve a variety of specific neurochemicals, including the peptide neurohormones oxytocin (OXY) and vasopressin (VP). OXY and VP derived from an evolutionary divergence of an ancestral hormone, vasotocin, that facilitates sociosexual responses in reptiles and birds (Moore, 1987). Recent evidence suggests that OXY and VP have special roles in social behavior, particularly nurturance and protective/territorial behavior. The following paragraphs summarize recent evidence implicating OXY and VP in prosocial behavior.

Prosociality in voles: The neurochemistry of monogamy and polygamy. An area of research that has produced some of the most remarkable findings involves voles: mouse-size rodents of the genus Microtus which exhibit both monogamous and polygamous lifestyles (Wang & Aragona, 2004). Monogamous prairie voles (Microtus ochrogaster) show lasting pair bonds that are activated by mating. When a female is exposed to a chemical in the urine of an unrelated male, she becomes sexually receptive, they mate repeatedly, and soon become parents. Bonded prairie voles live together in a common nest, snuggling side-by-side over 50% of the time. Either female or male will defend their territory by responding aggressively toward intruding voles. Both female and male care for their pups, with the male helping to build the nest and spending almost as much time with the young as the female; and if separated the pups become agitated and display ultrasonic distress calls and stress as evidenced by increases in cortisol. If the bonded partner dies, a surviving prairie vole will typically live alone rather than take a new mate. In contrast, the closely related meadow vole (Microtus pennsylvanicus) and montane vole (Microtus montanus) show none of the signs of strong social bonding displayed by the prairie vole. They are non-monogamous, nest independently, and breed promiscuously. The males play no parenting role, even the females abandon their pups soon after birth, and the pups do not appear to be distressed by abandonment (Carter, Lederhendler, & Kirkpatrick, 1997; Insel & Young, 2001). Insel (2002XX) has discussed this evidence in terms of a neurobiological basis for altruistic love, and implications for human altruism.

Oxytocin and VP are both critical to the social bonding of the prairie vole. Brain areas associated with OXY are larger in the prairie vole than the montane vole (Insel, Young, & Wang, 1997). In natural settings, mating is associated with OXY release, which

apparently functions to cement partner preferences in the female, so that she prefers the male she is with when OXY increase occurred (Carter, 1996). Experiments show that OXY is both necessary and sufficient for the development of pair bonds in the female: an unbonded female exposed to an unrelated male and OXY together will become bonded without mating, while an OXY antagonist will block bonding despite mating (Williams, Insel, Harbaugh & Carter, 1994).

Similarly, VP is necessary and sufficient for bonding in the male prairie vole: VP stimulates the formation of a partner preference even without mating, and VP antagonists prevent the formation of partner preferences even after extensive mating (Insel & Young, 2001). The distribution of VP receptors in the brains of monogamous and polygamous males is different, with the prairie vole having a higher density of VP receptors (termed V1aRs) in the ventral forebrain in an area associated with dopamine-mediated reward (Lim et al., 2004; Young, Lim, Gingrich, & Insel, 2001). Incredibly, the polygamous meadow vole male can be made to act like a monogamous prairie vole male by the alteration of a single gene: V1aR gene transfer, increasing VP receptors in the ventral forebrain of male meadow voles, substantially increased partner preference formation, indicating that “changes in the regional expression of a single gene can have a profound effect on the social behaviour of individuals within a species” (Lim et al. 2004, p. 756). The authors suggested that this has the effect of increasing the social memory of the partner’s olfactory signature, and associating it with dopamine-mediated reward: “monogamous social organization might be the result of the insertion of the V1aR system into the ancient pre-existing reward circuit” (p. 756).

Social recognition in mice. There is also evidence that prosocial neurochemical systems are involved in social behaviors in mice. OXY knock-out (OTKO) mice, bred without the genes to produce OXY, show profound deficits in social behavior (Young, 2001XX; 2002XX). As pups, OTKO mice are less distressed by social isolation, and they show no preference for their mothers compared to the strong preference shown by normal mice. As adults, OTKO mice demonstrate “social amnesia” in that they do not show evidence of learning to recognize other mice. This is reversed by a single injection of OXY into the brain prior to, but not after, exposure to another mouse. Young (2001XX) noted parallels between social deficits in OTKO mice and autism in humans: autism is characterized by profound social impairments, including deficits in social engagement and abnormal brain processing of social stimuli, and autistic children may have low concentrations of OXY in the blood.

Deprivation studies in monkeys and human beings. Although prosocial behavior is associated with specific neurochemical systems in the brain, early experience in social relationships appears to be critical for social attachment to develop normally. In Harry Harlow’s famous research, isolation for the first year of life rhesus monkeys was associated with serious socio-emotional deficits in adulthood (Harlow, 1971), and isolated animals manifested deficits in emotional communication as well (Miller, Caul, and Mirsky, 1967). These deficits are thought to be due to a lack of social experience at sensitive periods during development. In rhesus monkeys, strong attachment motives and emotions are present from birth, fearful and angry emotions develop after the first six months of age, and sexual emotions develop later at puberty. Harlow suggested that in the first three months, a Maternal (or Parental) Affectional System lays the socioemotional groundwork for the infant monkey’s emotional life, affording a basic

sense of trust in other monkeys; after six months, a Peer Affectional System allows the youngster to deal with and communicate fearful and angry emotions in the context of rough and tumble play; and these communication abilities are crucial in the last, Heterosexual Affectional System after the sexual systems come on line. Harlow (1971) suggested that these periods are analogous to infancy, childhood, and adolescence in human beings, and that if experience in earlier stages is lacking, it can have devastating effects on later social and emotional functioning.

Tragically, research with children who were institutionalized as infants in Eastern Europe have shown later socioemotional and emotional communication deficits that on the surface appear similar to those demonstrated in Harlow’s and Miller et al’s research (Pollak, Klorman, Thatcher, & Chicetti, 2001; Wismer Fries & Pollak, 2004). Recent evidence suggests that these children also show deficits in the OXY and VP systems (Wismer Fries, Ziegler, Kurian, Jacoris, & Pollak, 2005). These authors conclude that their results “suggest a potential mechanism whose atypical function may explain the pervasive social and emotional difficulties observed in many children who have experienced aberrant care-giving…consistent with the view that there is a critical role for early experience in the development of brain systems underlying basic aspects of human social behavior” (p. 17237).

The neurochemistry of trust. There is evidence that prosocial neurochemical systems my even be involved in the complex social deliberations associated with trust in human beings. Cooperation and competition have been studied in social dilemmas such as the Prisoner’s Dilemma (PD) game for decades. This research shows that participants often choose to cooperate, even where the payoffs in the situation would appear to encourage competition. It is noteworthy that one of the most powerful factors predicting cooperation is communication before the game (Sally, 1995). For example, Frank, Gilovich, and Regan (1993) allowed participants to interact prior to playing in a PD, and were allowed to converse about anything except the game. Cooperation was higher overall with communication was permitted, but interestingly the players seemed to accurately feel each other out: the dyads locked into either mutual cooperation or mutual defection, with relatively few cases of one partner taking advantage of the other. The intention to cooperate or defect was apparently communicated prior to play. Boone and Buck (2003) suggested that emotional expressivity acts as a marker for trustworthiness, and allows trust or suspicion to be displayed nonverbally; and Rauh, Polonsky, and Buck (2004) found friendly emotional expressiveness and suspiciousness to predict a participant’s cooperative or competitive first move, respectively, in a PD game.

Recent evidence suggests that OXY may play a role in cooperation in human beings. Kosfeld et al. (2005) found that OXY presented in an aerosol increased cooperative behavior, and Wood et al (2005) found that disruption of serotonin functioning increases competitive behavior in PD games. Thus, remarkably, molecules relevant to attachment mechanisms can have significant effects upon complex human behavior. It thus appears that primordial affective motivational-emotional systems can influence complex choices in human beings involving cooperation and competition.

Other Prosocial Neurochemical SystemsAlthough evidence for the prosocial functions of OXY and VP is particularly

impressive, other neurochemical systems have prosocial functions as well (See Buck,

1999; Panksepp, 1998 for reviews). These include the endogenous opiates or endorphins, which Panksepp and colleagues have associated with playful behavior in rats (Panksepp & Burgdorf, 1999; 2000). Burgdorf and Panksepp (2001) suggest that the positively reinforcing effects of play are modulated by the endorphins, fitting with evidence of the role of endorphins in mediating social reward (Panksepp, 1991). Also, gonadotropin-releasing hormone (GnRH) increases sexual proclivities and has rewarding effects as demonstrated by its ability to establish conditioned place preferences in male rats: these preferences, however, are absent in gonadectomized animals (deBeun et al., 1991). Panksepp (1993) has suggested that GnRH “may well prove to be a prime mover in human libido” (p. 94).

Attachment and Higher-Level Social and Moral Emotions

I have argued that cooperative as well as competitive tendencies have been built-into the genome from the beginning, and that specific neurochemical systems in the brain are associated with powerful prosocial motives and emotions. These powerful prosocial emotions are hiding in plain sight: while they are involved in motivating much of the behavior of interest to social psychology, they tend to be taken for granted and their importance is rarely recognized. Just as effectance motives and needs for understanding underlie tendencies toward cognitive consistency, attribution processes, and attitude formation and change; attachment motives and needs to be loved underlie tendencies toward social referencing, modeling and imitation, conformity, and obedience (Buck, 1998). They are involved in every human action and interaction.

Defining Motivation and EmotionDefinitions of “motivation” generally emphasize the activation and direction of

behavior toward a goal; and definitions of “emotion” also involve goal-directed action but typically note the presence of peripheral physiological responses, expressive behaviors and subjective experiences or affects (Kleinginna and Kleinginna 1981a, 1981b). Developmental-interactionist (DI) theory suggests that motives and emotions imply one another: that motives constitute potential for behavior built into a system of behavior control, and that emotions constitute readouts of motivational potential when that potential is activated by an effective stimulus. There are three sorts of readout: Emotion I involves peripheral physiological responding via the autonomic, endocrine, and immune systems; Emotion II involves expressive displays such as facial expressions and pheromone releases; and Emotion III involves subjective or affective experience of desires and feelings (Buck, 1988a). In this view, motivation and emotion are related similarly to energy and matter in physics. Energy is a potential that is never seen in itself, but only in its manifestation in matter when activated by an effective stimulus: in heat, light, and/or force. Thus, the potential energy in an explosive or a coiled spring is not observable in itself, but it is revealed when activated by lighting the fuse or releasing the spring. Similarly, motivation is never seen in itself, but only in its manifestations in emotion: that is, in goal-directed behavior, peripheral physiological responding, expressive displays, and subjectively experienced desires and feelings.

One feature of the DI view is that it implies that all of the biologically-based motives and emotions are always turned on. Neurochemical systems underlying the experience

and expression of happiness, sadness, fear, anger, sex, hunger, thirst, etc. are always activated to some extent, and we can turn our attention to the experience of these emotions. But, like the feel of the shoes on our feet, this activation is typically weak and these feeling are unnoticed unless an effective stimulus is presented.

Evolutionary Roots of Motivation and EmotionPrimes and paleoprimes. The above definitions can encompass primary

motivational/emotional states (primes) in human beings and also animals, with the complexity of the state compatible with the complexity of the brain of the species in question. DI theory argues that the capacity for the subjective experience of primary emotions such as happiness, sadness, fear, and anger are associated with activity in specific neurochemical systems, and studies of drug discrimination and place learning suggest that animals are also capable of “subjective experience” (Buck, 1999; Panksepp, 1998). However, these definitions can also apply to primary motivational-emotional systems in microbes whose evolution long preceded the evolution of the brain, which one might term paleoprimes. Paleoprimes serve functions in simple creatures analogous to those served by emotions in human beings—activation, approach-avoidance, dispersion, and aggregation—and these can be observed in their behavior (Buck, 1999). Microbes are active at some times and quiescent at others. They can be observed to approach some stimuli (light, warmth) and avoid others. And, as we have seen, microbes also achieve dispersion, aggregation, and cooperation by releasing molecules (pheromones) that repel, attract, or otherwise influence others of their species.

Paleosociality. These systems in turn underlie paleosociality, involving the abilities of microbes to communicate with one another and thereby complete social structures and organizations (Buck & Renfro, 2006). As communication occurs in the course of interaction between individual microbes, social organization emerges spontaneously and effortlessly as a self-organizing system. The mechanism of this emergence through interaction is spontaneous communication between elements, that is, between individual microbes. In these creatures, the systems are simple enough that specific sending and receiving mechanisms can be specified and the system of emergence understood at a molecular and even genetic level. I suggest that these microbial systems illustrate fundamental principles of social organization that illuminate mechanisms and principles of social organization in more complex creatures including human beings.

More specifically, I suggest that a combination of attachment motives and social comparison processes underlie the natural and effortless emergence of systems of social and moral emotions that guide every human interchange. Motivational potential inherent in attachment systems in infants are activated through early interactions with caregivers, giving rise to fundamental needs to love and to be loved which constitute the affective foundation of sociality. The need to be loved is, therefore, fundamental to human sociality. With development, interactions with caregivers and peers teach the child rules that must be followed in order to be loved: thus the child learns that one must meet and exceed expectations in order to be loved. The need to meet and exceed expectations therefore requires both a need to be loved and learning what those rules and expectations are and how to meet them. This learning occurs naturally, effortlessly, and largely unconsciously during the course of interactions with caregivers, peers, and later, lovers.

Primary Social and Moral EmotionsUnlike biologically-based emotions, DI theory regards social, moral, and also

cognitive emotions to be higher-level emotions, requiring both a physiological base in neurochemical systems associated with attachment and exploration, and a “cognitive” consideration of situational and interpersonal contingencies (Buck, 1999). The neurochemical systems provide the affective “fire” to the higher-level emotions, while the contingencies determine the quality of the emotion. More specifically, the individual person (P) is exquisitely aware of success or failure in meeting expectations and being loved, and P’s own success or failure can be compared with the success or failure of comparison others (O’s). All possible combinations of success and failure of self and other yields eight combinations of fundamental interpersonal contingencies that correspond to eight primary social emotions (PSEs) organized into four pairs of twin social emotions: if one succeeds one experiences pride/arrogance, if one fails it is guilt/shame, if the other succeeds it is envy/jealousy, if the other fails it is pity/scorn. The first of these twins is associated with meeting/exceeding expectations, that second with being loved.

Attachment and social emotions. There is much evidence in the extensive research stemming from attachment theory (Bowlby, 1969/1982) that persons who are securely attached are relatively certain of being loved, while persons with attachment anxiety worry that they are not loved, and persons with avoidant attachment distrust others and in effect eschew love (e.g., Mikulciner, 1998; Mikulciner & Shaver, 2003, 2005; Mikulciner, Shaver, Gillath, & Nitzberg, 2005). I suggest that secure attachment therefore is associated with relatively less attention to being loved and greater attention to meeting/exceeding expectations; that anxious attachment is associated with a greater attention to being loved; and avoidant attachment is associated with relatively low prosocial needs and therefore relatively weak social emotions. Therefore, a secure person would be expected to experience pride, guilt, envy, and pity in situations where an anxious person would experience arrogance, shame, jealousy, and scorn; and that an avoidant person would tend to not experience any of these emotions strongly with the extreme being a psychopath with no need to be loved who is incapable of experiencing any of the social or moral emotions.

In this analysis, attachment is viewed as both a trait and a state. Most children can be classified as having a specific attachment style, while most adults show a mixed style. This may be because as social development proceeds, attachment security can increasingly vary with the personal relationship of P vis a vis O: we may be secure that we are loved by some persons but anxious about the love of others. So, we tend to feel pride in comparison with the former and arrogance in comparison with the latter, for example. Also, we can learn to avoid attachment with some persons. It is all too easy to learn that certain persons are enemies who do not deserve our love, to whom social and moral emotions are irrelevant. The ability of even normal human beings to become situational psychopaths—to destroy others without moral compunction—is all too apparent, as Hannah Arndt showed in her analysis of the banality of evil (Arndt, 1951/1973).

The dynamics of social emotions. This analysis implies that the eight PSEs are interrelated: that for example when a person is proud they tend to pity others, and to NOT to feel guilt or envy of others. Makoto Nakamura, Edward Vieira, Maxim Polonsky, and

I have tested this by giving scenarios about comparative success and failure to University students in the United States and Japan: for example, if another person won the lottery and got rich, how would you feel about them, and how would they feel about you? This study found support for the hypothesized relationships between primary social emotions in virtually every case, whether labeled in English or Japanese. This supported the hypothesis of universal labeling: that the words for the eight PSEs could be found in all languages, and the hypothesis of universal dynamics: that they would be interrelated similarly in all languages. There were interesting differences between Japan and the United States as well, consistent with the collectivist-individualist distinction: Japanese were consistently more embarrassed than Americans, and Americans more proud than Japanese, across all of the scenarios. It was also interesting that in both the US and Japan, pity and scorn were negatively correlated, suggesting that there are two ways of responding to the less fortunate, perhaps related to the security of the individual: secure persons feel pity while anxious person feel scorn when the other fails (Buck, Nakamura, Vieira, and Polonsky, 2005, submitted for publication).

Primary moral emotions. This analysis has been extended to the analysis of eight primary moral emotions (PMEs), where success or failure for self and other is combined with the judgment that the outcome is just or unjust. When one’s success is just the result is triumph, when unjust it is modesty; when one’s failure is just the result is humiliation, when unjust it is indignation; when the other’s success is just the result is admiration, when unjust it is resentment; when the other’s failure is just the result is contempt, when unjust it is sympathy. The PMEs are related to the PSEs: for example, envy and jealousy can go with either admiration or resentment depending upon whether the other's success is seen as justified. Research is presently under way to analyze the labeling and dynamics of PSEs and PMEs in Japan, Ukraine/Russia, and India with Makoto Nakamura, Maxim Polonsky and Sripriya Rangarajan.

Dominance-submission versus civility. The dynamics of social and moral emotions are illustrated in Figure 1, which illustrates the social and moral emotions in a situation of dominance and submission. The successful figure on the left exudes triumph, pride and arrogance; and regards the inept figure on the right with contempt and scorn. The unsuccessful figure feels guilt, shame, humiliation, and indignation; and regards the other with envy, jealousy, and resentment. These can be strong acute emotions; they can be fleeting as one might feel as one sees a person driving by in an expensive car, or they can be chronic, unremitting, and grinding; leading to a lack of authentic social contact and communication, the exacerbation of stress, unhappiness, and depression—perhaps in both parties—whose true cause may be unrecognized.

---Figure 1---On the other hand, if the two figures show each other civility, a pattern of positive

interpersonal emotions can result. Even despite differences in wealth and status, it is possible for two persons to have a relationship of mutual trust and respect in which each regard the other as following the rules fairly and with a sense of justice. Each can then regard their own successes with modesty, and respond to the other with a sense of mutual gratitude for following the rules and admiration for their deserve success. These are the ingredients for authentic communication and the powerful stress-buffering that can come with social support.

This view of PSE's and PME's implies that sociality and morality arise spontaneously and effortlessly from interaction: a spontaneous restructuring of socio-emotional experience analogous to Piaget's (1971) assimilation and accommodation process in the realm of cognitive development. From this point of view, organized religion is not necessary to morality, and indeed can potentially get in the way of morality: even promoting the situational psychopathy of religious intolerance and conflict. Similarly, as Benson Ginsburg and I have suggested, this viewpoint suggests that kin selection and reciprocity, rather than being the bases of altruism, are actually mechanisms to restrict loving and altruistic feelings to kin and comrade: rather than underlying altruism, kin selection and reciprocity are at the roots of xenophobia.

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Figure Captions

Figure 1. In a dominant-submissive relationship, conflict can be exacerbated by moral considerations: The perception of unfairness.