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Altruism: its characteristics and evolution.

Altruism is a group phenomenon in which some genes or individuals, which must be presumed to be selfish, benefit others at cost to themselves. The presumption of selfishness and the fact of altruism are reconciled by kin-group selection and by reciprocal altruism. Kin-group selection is clearly visible only in special cases; its role even among social insects may be overestimated; it is probably usually inhibited by competition. However, reciprocal altruism is ubiquitous. All altruism is: (i) potentially reciprocal; (ii) potentially profitable to altruists as well as to recipients; (iii) environmentally determined, usually by position of individuals in group or environmental situations; and (iv) a net-gain lottery. These generalizations are illustrated by four idealized cases; the difficulty of applying them to real cases is illustrated by alarm-calling in groups of birds. Although altruism is a group phenomenon, it evolves by individual selection, by processes equivalent to co-evolutions. Its evolution is: (i) opposed by competition; (ii) costly, complex, and slow, and tending to produce an imprecise flexible altruism rather than a precisely detailed one; and (iii) supplemented by group selection (differential extinction of groups). That altruism in human beings conforms to these generalizations is a good working hypothesis. However, analysis does not "take the altruism out of (human) altruism." Humans do not calculate it, but behave altruistically because they have human altruistic emotions.

Altruism

Socialization and situational influences on sustained altruism.

In this conceptual replication and extension of Rosenhan's study of civil rights activists, the sustained altruism (i.e., help that extends over time) of volunteers at a telephone crisis-counseling agency was examined. Using a prospective format, it was predicted that volunteers with a socialization history of exposure to nurturant parents who modeled altruism (autonomous altruists) would exhibit a greater degree of sustained altruism than those with a history of less nurturant parents who modeled altruism to a lesser degree (normative altruists). The altruism of the normative volunteers, however, was expected to increase given certain situational conditions (here, participation in a highly cohesive training group prior to the actual volunteer activity). As predicted, the rate of sustained altruism of normative volunteers in highly cohesive groups was increased to a level comparable to that of autonomous volunteers, while the altruism of autonomous volunteers was not affected by the training group experience. The implications of these findings for research on altruism and its development, as well as some applications to volunteerism, are discussed.

Adolescent

Gene-culture coevolution: models for the evolution of altruism with cultural transmission.

Models of sexual haploids under kin selection are constructed. The trait of altruism is transmitted vertically from parent to child, but not in a strictly genetic manner. Two systems of altruism are considered: parent-to-offspring and sib-to-sib. In the former case it is shown that even when Hamilton's conditions for the success of genetically determined altruism are met, genes that increase the transmission of altruism may not invade the population. With sib-to-sib altruism, such genes will always increase initially.

Alleles

A mechanism for social selection and successful altruism.

Within the framework of neo-Darwinism, with its focus on fitness, it has been hard to account for altruism behavior that reduces the fitness of the altruist but increases average fitness in society. Many population biologists argue that, except for altruism to close relatives, human behavior that appears to be altruistic amounts to reciprocal altruism, behavior undertaken with an expectation of reciprocation, hence incurring no net cost to fitness. Herein is proposed a simple and robust mechanism, based on human docility and bounded rationality that can account for the evolutionary success of genuinely altruistic behavior. Because docility-receptivity to social influence-contributes greatly to fitness in the human species, it will be positively selected. As a consequence, society can impose a "tax" on the gross benefits gained by individuals from docility by inducing docile individuals to engage in altruistic behaviors. Limits on rationality in the face of environmental complexity prevent the individual from avoiding this "tax." An upper bound is imposed on altruism by the condition that there must remain a net fitness advantage for docile behavior after the cost to the individual of altruism has been deducted.

Altruism

"Runaway" social evolution: reinforcing selection for inbreeding and altruism.

Kin selection theory predicts that altruistic behaviors, those that decrease the fitness of the individual performing the behavior but increase the fitness of the recipient, can increase in frequency if the individuals interacting are closely related. Several studies have shown that inbreeding therefore generally increases the effectiveness of kin selection when fitnesses are linear, additive functions of the number of altruists in the family, although with extreme forms of altruism, inbreeding can actually retard the evolution of altruism. These models assume that a constant proportion of the population mates at random and a constant proportion practices some form of inbreeding. In order to investigate the effect of inbreeding on the evolution of altruistic behavior when the mating structure is allowed to evolve, we examined a two-locus model by computer simulation of a diploid case and illustrated the important qualitative features by mathematical analysis of a haploid case. One locus determines an individual's propensity to perform altruistic social behavior and the second locus determines the probability that an individual will mate within its sibship. We assumed positive selection for altruism and no direct selection at the inbreeding locus. We observed that the altruistic allele and the inbreeding allele become positively associated, even when the initial conditions of the model assume independence between these loci. This linkage disequilibrium becomes established, because the altruistic allele increases more rapidly in the inbreeding segment of the population. This association subsequently results in indirect selection on the inbreeding locus. However, the dynamics of this model go beyond a simple "hitch-hiking" effect, because high levels of altruism lead to increased inbreeding, and high degrees of inbreeding accelerate the rate of change of the altruistic allele in the entire population. Thus, the dynamics of this model are similar to those of "runaway" sexual selection, with gene frequency change at the two loci interactively causing rapid evolutionary change.

Alleles

[The relation between the attitude toward humor, aggression and altruism].

This study investigated the attitude towards humor and the relationship among the attitude, aggression and altruism. Four hundred and seven subjects answered the questionnaires consisting of (1) items about attitude toward humor, (2) items about aggression and altruism, and (3) items about humorous statements, frequently heared in negative situations. Factor analysis revealed that the attitude toward humor consists of two factors; (a) preference for aggressive humor and (b) preference for playful humor. The former is related to aggression and altruism, while the latter is related to altruism only. The items about humorous statements frequently heard in negative situations suggested that preference for aggressive humor is related to aggressive behavior, while preference for playful humor is related to moderation of the atmosphere.

Adult

Nonmathematical models for evolution of altruism, and for group selection (peck order-territoriality-ant colony-dual-determinant model-tri-determinant model).

Mathematical biologists have failed to produce a satisfactory general model for evolution of altruism, i.e., of behaviors by which "altruists" benefit other individuals but not themselves; kin selection does not seem to be a sufficient explanation of nonreciprocal altruism. Nonmathematical (but mathematically acceptable) models are now proposed for evolution of negative altruism in dual-determinant and of positive altruism in tri-determinant systems. Peck orders, territorial systems, and an ant society are analyzed as examples. In all models, evolution is primarily by individual selection, probably supplemented by group selection. Group selection is differential extinction of populations. It can act only on populations preformed by selection at the individual level, but can either cancel individual selective trends (effecting evolutionary homeostasis) or supplement them; its supplementary effect is probably increasingly important in the evolution of increasingly organized populations.

Animals

Genetics of mutualism: the evolution of altruism between species.

Conditions are analyzed under which natural selection favors an individual to help another species at a cost to its own reproduction. Traditional models for the evolution of altruism between species focus on the genetic relatedness between the original donor and the recipients of return benefits from the mutualistic partner species. A more general model is analyzed here that focuses on the synergistic effects between partner species caused by genetic variability. The model shows that the spread of altruism is enhanced by spatial correlations between species in the genetic tendency to give aid to partners. These spatial correlations between species are similar to the kin selection coefficients of relatedness that determine the course of social evolution within species. The model also shows that natural selection and ecological dynamics can create genetic correlations between neighbors of different species, even when the initial spatial distributions of the species are uncorrelated. Genetic correlations between species may play an important role in the origin and maintenance of altruism between species.

Altruism

Altruism and aggression: the heritability of individual differences.

Five questionnaires measuring altruistic and aggressive tendencies were completed by 573 adult twin pairs of both sexes from the University of London Institute of Psychiatry Volunteer Twin Register. The questionnaires measured altruism, empathy, nurturance, aggressiveness, and assertiveness. The intraclass correlations for the five scales, respectively, were .53, .54, .49, .40, and .52 for 296 monozygotic pairs, and .25, .20, .14, .04, and .20 for 179 same-sex dizygotic pairs, resulting in broad heritability estimates of 56%, 68%, 70%, 72%, and 64%. Additional analyses, using maximum-likelihood model-fitting, revealed approximately 50% of the variance on each scale to be associated with genetic effects, virtually 0% with the twins' common environment, and the remaining 50% with each twins' specific environment and/or error associated with the test. Correcting for the unreliability in the tests raised the maximum-likelihood heritabilities to approximately 60%. Age and sex differences were also found: altruism increased over the age span from 19 to 60, whereas aggressiveness decreased, and, at each age, women had higher scores than men on altruism and lower scores on aggressiveness.

Adult

An analysis of altruism: a concept of caring.

In clinical settings, nurses have observed a conflict within women who are expected to focus on caring for themselves, while their value lies in attending to the needs of others at the sacrifice of self. In this concept analysis of altruism, the term is defined and the critical attributes are identified. Model, borderline, related and contrary cases illustrate what the concept is and what it is not. The meaning of altruism is compared to self-neglect and co-dependence, which are two similar concepts. The antecedents and consequences of altruism are identified to further refine the critical attributes. Implications for research are discussed. This is a first step towards developing nursing theory that will give direction for providing more effective care to women within their value system.

Altruism

Inclusive fitness, altruism and family adaptation.

An integrative model of family functioning is put forward using a sociobiological framework. Three key sociobiological concepts that have a relevance to family interaction are inclusive fitness, altruism, and a struggle for status. The term 'inclusive fitness' encompasses the parents' own fitness and that of their kith and kin. Altruism refers to how parents promote the survival of their progeny and extended family. The struggle for status refers to a struggle for power and prestige. According to the author's hypothesis 'inclusive fitness' exerts its influence through the struggle for status which in turn influences the degree of altruism and the level of 'family adaptation'. The parents' level of inclusive fitness largely determines the level of family adaptation which in turn influences the growth and individuation of the next generation. A common negative outcome of failure in the struggle for status and power is a displacement of the struggle for power into a family context. Failure in a struggle for status between peer group members would in a pretechnological society probably have reflected a relative lack of fitness. Failure in the struggle for status is likely to generate maladaptive cycles in the family which will lower the inclusive fitness of the parents. A concept of ideal family functioning is derived from sociobiology and specifically related to a high inclusive fitness in the parents. The author argues that sociobiological concepts can contribute to the development of an integrative theory of family behaviour. The clinical implications of this model are explored.

Adaptation, Psychological

Non-patient decision-making in medicine: the eclipse of altruism.

Despite its virtues, lay decision-making in medicine shares with professional decision-making a disturbing common feature, reflected both in formal policies prohibiting high-risk research and in informal policies favoring treatment decisions made when a crisis or change of status occurs, often late in a downhill course. By discouraging patient decision-making but requiring dedication to the patient's interests by those who make decisions on the patient's behalf, such practices tend to preclude altruistic choice on the part of the patient. This eclipse is to be regretted not just because widescale altruism has the capacity to provide important social goods and correct injustices in distribution, but for intrinsic reasons as well. It is argued that preserving the possibility of altruism obliges patients--and future patients--to make decisions about dying and other medical matters in advance, thus avoiding that displacement of decision-making onto lay and professional second parties which results in altruism's eclipse.

Altruism

Altruism as hedonism: a social development perspective on the relationship of negative mood state and helping.

A study was conducted to provide a means for reconciliation of the conflicting data on the relationship of negative mood state to altruism. Whereas some studies have shown that negative mood leads to increases in altruistic action, others have shown the reverse. It was hypothesized that the inconsistency of these results was due to differences in the ages and consequent levels of socialization of the subjects employed in the earlier studies. In order to test the hypothesis, subjects from three age groups (6-8, 10-12, and 15-18 years old) were asked to think of either depressing or neutral events and were subsequently given the opportunity to be privately generous. Consistent with predictions from the negative state relief model of altruism, the youngest, least socialized subjects were somewhat less generous in the negative mood condition, but this relationship progressively reversed itself until in the oldest, most socialized group, the negative mood subjects were significantly more generous than neutral mood controls. The data were taken as support for a hedonistic conception of altruism that views adult benevolence as self-gratification. It is suggested that the reward character of benevolence derives from the socialization experience.

Adolescent

Altruism and the practice of medicine.

Biological and psychological research into the antecedents of altruism has considerable significance for those involved in the teaching or practice of medicine. The evidence available indicates that altruistic behaviour is a universal phenomenon influenced by intra-individual, interpersonal, situational and sociocultural factors. A central theme which emerges is the strikingly ambivalent nature of the altruistic drive. The factors which may facilitate or inhibit altruism in medical students and doctors are discussed. Some ways of ensuring that medical training does not seriously distort altruism are suggested.

Achievement

Critical self-reflection and self-perceived altruism: when self-reward fails.

Social learning theory has produced a three-step model of prosocial development: In the young child, prosocial behavior is elicited by material rewards; in the older child, it is elicited also by social rewards; and in the morally mature adult, it is elicited by self-rewards based on an internalized perception of oneself as a kind, caring, altruistic individual. Self-perception theory has complicated this social learning model by demonstrating that once the third step is reached, the continued presence of material and social rewards may undermine intrinsic prosocial motivation based on self-rewards, producing moral regression. We propose a further complication. Critical self-reflection--the desire to know thyself, warts and all--may introduce a self-deprecating attributional bias that can undermine self-perceived altruism, even following helping for which one receives only intrinsic self-rewards. Two experiments are reported in which we manipulated critical self-reflection on one's reasons for helping. Results indicated that self-reflection undermined self-perceived altruism, especially when the salience of the self-rewards for helping was high. Experiment 2 also provided evidence that, as predicted, this self-reflection effect was most apparent for individuals who valued self-knowledge more highly than concern for others. Moral consequences of critical self-reflection are discussed.

Altruism

Five studies testing two new egoistic alternatives to the empathy-altruism hypothesis.

The empathy-altruism hypothesis claims that prosocial motivation associated with feeling empathy for a person in need is directed toward the ultimate goal of benefiting that person, not toward some subtle form of self-benefit. We explored two new egoistic alternatives to this hypothesis. The empathy-specific reward hypothesis proposes that the prosocial motivation associated with empathy is directed toward the goal of obtaining social or self-rewards (i.e., praise, honor, and pride). The empathy-specific punishment hypothesis proposes that this motivation is directed toward the goal of avoiding social or self-punishments (i.e., censure, guilt, and shame). Study 1 provided an initial test of the empathy-specific reward hypothesis. Studies 2 through 4 used three procedures to test the empathy-specific punishment hypothesis. In Study 5, a Stroop procedure was used to assess the role of reward-relevant, punishment-relevant, and victim-relevant cognitions in mediating the empathy-helping relationship. Results of these five studies did not support either the empathy-specific reward or the empathy-specific punishment hypothesis. Instead, results of each supported the empathy-altruism hypothesis. Evidence that empathic emotion evokes altruistic motivation continues to mount.

Adult

Empathic joy and the empathy-altruism hypothesis.

Three experiments tested whether empathy evokes egoistic motivation to share vicariously in the victim's joy at improvement (the empathic-joy hypothesis) instead of altruistic motivation to increase the victim's welfare (the empathy-altruism hypothesis). In Experiment 1, Ss induced to feel either low or high empathy for a young woman in need were given a chance to help her. Some believed that if they helped they would receive feedback about her improvement; others did not. In Experiments 2 and 3, Ss induced to feel either low or high empathy were given a choice of getting update information about a needy person's condition. Before choosing, they were told the likelihood of the person's condition having improved--and of their experiencing empathic joy--was 20%, was 50%, or was 80%. Results of none of the experiments patterned as predicted by the empathic-joy hypothesis; instead, results of each were consistent with the empathy-altruism hypothesis.

Adult

Grooming, alliances and reciprocal altruism in vervet monkeys.

Reciprocal altruism refers to the exchange of beneficial acts between individuals, in which the benefits to the recipient exceed the cost to the altruist. Theory predicts that cooperation among unrelated animals can occur whenever individuals encounter each other regularly and are capable of adjusting their cooperative behaviour according to experience. Although the potential for reciprocal altruism exists in many animal societies, most interactions occur between closely related individuals, and examples of reciprocity among non-kin are rare. The field experiments on vervet monkeys which we present here demonstrate that grooming between unrelated individuals increases the probability that they will subsequently attend to each others' solicitations for aid. Vervets appear to be more willing to aid unrelated individuals if those individuals have behaved affinitively toward them in the recent past. In contrast, recent grooming between close genetic relatives appears to have no effect on their willingness to respond to each other's solicitations for aid.

Altruism