Aggressive contests probably occur in networking environments where information about fighting ability is conveyed both to an opponent and to individuals peripheral to the fight itself, the bystanders. Our primary aim was to investigate the relative influences of eavesdropping and prior social experience on the dynamics of aggressive contests in Xiphophorus helleri. A bystander's ability to witness an encounter was manipulated using clear, one-way mirror, and opaque partitions. After watching (or not watching) the initial contest, the bystander encountered either the winner or loser of the bout. Treatment comparisons of bystander-winner or bystander-loser contest dynamics indicated the presence or absence of winner, loser, or eavesdropping effects. Winner and loser effects had negligible influences on bystander contest dynamics. Eavesdropping significantly reduced the bystander's propensity to initiate aggression, escalate, and win against seen winners regardless of whether the watched bout had escalated or not. Though eavesdropping had relatively little effect on bystander-loser contest dynamics, bystanders were less prone to initiate aggression and win against losers that had escalated in the witnessed bout. Thus, bystanders appear to preferentially retain and utilize information gained about potentially dangerous opponents (winners or persistent losers). Our data lend clear support for the importance of eavesdropping in visually based aggressive signalling systems.
Winner and loser effects, in which the outcome of an aggressive encounter influences the tendency to escalate future conflicts, have been documented in many taxa, but we have limited understanding of why they have evolved. One possibility is that individuals use previous victories and defeats to assess their fighting ability relative to others. We explored this idea by modelling a population of strong and weak individuals that do not know their own strength, but keep track of how many fights they have won. Under these conditions, adaptive behaviour generates clear winner and loser effects: individuals who win fights should escalate subsequent conflicts, whereas those who lose should retreat from aggressive opponents. But these effects depend strongly on age and experience. Young, naive individuals should show highly aggressive behaviour and pronounced loser effects. For these inexperienced individuals, fighting is especially profitable because it yields valuable information about their strength. Aggression should then decline as an individual ages and gains experience, with those who lose fights becoming more submissive. Older individuals, who have a better idea of their own strength, should be more strongly influenced by victories than losses. In conclusion, we predict that both aggressiveness and the relative magnitude of winner and loser effects should change with age, owing to changes in how individuals perceive their own strength.
winner effect; loser effect; state-dependent aggression; dynamic model; Hawk–Dove game
Animal conflicts are influenced by social experience such that a previous winning experience increases the probability of winning the next agonistic interaction, whereas a previous losing experience has the opposite effect. Since androgens respond to social interactions, increasing in winners and decreasing in losers, we hypothesized that socially induced transient changes in androgen levels could be a causal mediator of winner/loser effects. To test this hypothesis, we staged fights between dyads of size-matched males of the Mozambique tilapia (Oreochromis mossambicus). After the first contest, winners were treated with the anti-androgen cyproterone acetate and losers were supplemented with 11-ketotestosterone. Two hours after the end of the first fight, two contests were staged simultaneously between the winner of the first fight and a naive male and between the loser of first fight and another naive male. The majority (88%) of control winners also won the second interaction, whereas the majority of control losers (87%) lost their second fight, thus confirming the presence of winner/loser effects in this species. As predicted, the success of anti-androgen-treated winners in the second fight decreased significantly to chance levels (44%), but the success of androgenized losers (19%) did not show a significant increase. In summary, the treatment with anti-androgen blocks the winner effect, whereas androgen administration fails to reverse the loser effect, suggesting an involvement of androgens on the winner but not on the loser effect.
social experience; winner effect; androgens; testosterone; aggression
Many animals use information acquired from recent experiences to modify their responses to new situations. Animals’ decisions in contests also depend on their previous experience: after recent victories individuals tend to behave more aggressively and after defeats more submissively. Although these winner and/or loser effects have been reported for animals of different taxa, they have only recently been shown to be flexible traits, which can be influenced by extrinsic factors. In a mangrove killifish (Kryptolebias marmoratus), for instance, individuals which lost an earlier contest were more likely than others to alter contest decisions after a recent win/loss. This result suggests that individuals perceiving themselves to have worse fighting abilities are more inclined to adjust contest strategy based on new information. If this is the case, an individual’s propensity to modify behaviour after a win/loss might also be modulated by intrinsic mechanisms related to its ability to fight. Stress and sex steroid hormones are often associated with an individual’s contest behaviour and performance, so, in this study, we tested the hypothesis that an individual’s propensity to change behaviour after wins or losses also depends on its hormonal state.
Our results show that an individual’s propensity to adjust contest decisions after wins and losses does depend on its hormonal state: individuals with lower levels of cortisol (F), testosterone (T) and 11-ketotestosterone (KT) are more receptive than others to the influence of recent contest experiences, especially losing experiences, and the influences last longer. Furthermore, although winning and losing experiences resulted in significant changes in behaviour, they did not bring about a significant change in the levels of F, T, KT or oestradiol (E2).
This study shows that an individual’s receptivity to the influence of recent wins and losses is modulated by its internal state, as well as by extrinsic factors. Individuals with hormonal profiles corresponding to lower aggressiveness and a reduced likelihood of winning were more likely to alter contest decisions after a recent win/loss. The results also suggest that F, T, KT and E2 are not the primary physiological mechanisms mediating winner-loser effects in this fish.
Animal contest; Information; Winner-loser effect; Cortisol; 11-ketotestosterone; Testosterone; Oestradiol; Kryptolebias marmoratus
In the field, phenotypic determinants of competitive success are not always absolute. For example, contest experience may alter future competitive performance. As future contests are not determined solely on phenotypic attributes, prior experience could also potentially alter phenotype–fitness associations. In this study, we examined the influence of single and multiple experiences on contest outcomes in the jumping spider Phidippus clarus. We also examined whether phenotype–fitness associations altered as individuals gained more experience. Using both size-matched contests and a tournament design, we found that both winning and losing experience affected future contest success; males with prior winning experience were more likely to win subsequent contests. Although experience was a significant determinant of success in future contests, male weight was approximately 1.3 times more important than experience in predicting contest outcomes. Despite the importance of experience in determining contest outcomes, patterns of selection did not change between rounds. Overall, our results show that experience can be an important determinant in contest outcomes, even in short-lived invertebrates, and that experience alone is unlikely to alter phenotype–fitness associations.
jumping spider; multiple competition; Phidippus clarus; previous experience; selection gradient; tournament design
Coalition formation has been documented in a diverse array of taxa, yet there has been little formal analysis of polyadic interactions such as coalitions. Here, we develop an optimality model which examines the role of winner and loser effects in shaping coalition formation. We demonstrate that the predicted patterns of alliances are strongly dependent on the way in which winner and loser effects change with contestant strength. When winner and loser effects decrease with the resource-holding power (RHP) of the combatants, coalitions will be favoured between the strongest members of a group, but not between the weakest. If, in contrast, winner and loser effects increase with RHP, exactly the opposite predictions emerge. All other things being equal, intervention is more likely to prove worthwhile when the beneficiary of the aid is weaker (and its opponent is stronger), because the beneficiary is then less likely to win without help. Consequently, intervention is more probable when the impact of victory on the subsequent performance of a combatant increases with that individual's strength because this selects for intervention in favour of weaker combatants. The published literature on hierarchy formation does not reveal how winner and loser effects actually change with contestant strength and we therefore hope that our model will spur others to collect such data; in this light we suggest an experiment which will help to elucidate the nature of winner and loser effects and their impact on coalition formation in animals.
Agonistic behaviour between male orb-web spiders Metellina mengei competing for access to female webs was examined in field experiments to test the major predictions of game theory. Winners of fights were significantly larger than losers, particularly with respect to the length of the first pair of legs, which are sexually dimorphic in this species and used extensively in agonistic encounters. The size of the winning male had no influence on contest intensity or duration, and neither did relative size. However, fight intensity and duration were both positively correlated with the size of the losing male. Resident males won significantly more contests than intruders. Winning intruders were significantly larger than winning residents and it was these winning intruders that tended to produce the longer fights. Female weight and hence reproductive value had a marked influence on fight intensity and duration of fights won by the intruder but not those won by the resident. This indicates that only the resident obtains information about the female. These data are discussed with reference to the discrepancy with theory and a failure of some contestants to obtain information on resource value and relative contestant size necessary to optimize fight strategy.
Contest theory predicts the evolution of a stable mixture of different strategies for fighting. Here, we investigate the possibility that stable between-individual differences in startle-response durations influence fighting ability or ‘resource-holding potential’ (RHP) in the beadlet sea anemone, Actinia equina. Both winners and losers showed significant repeatability of pre-fight startle-response durations but mean pre-fight startle-response durations were greater for eventual losers than for eventual winners, indicating that RHP varies with boldness. In particular, individuals with short startle responses inflicted more attacks on their opponent. Both repeatability and mean-level responses were changed by the experience of fighting, and these changes varied with outcome. In losers, repeatability was disrupted to a greater extent and the mean startle-response durations were subject to a greater increase than in winners. Thus, following a fight, this behavioural correlate of RHP behaves in a way similar to post-fight changes in physiological status, which can also vary between winners and losers. Understanding the links between aggression and boldness therefore has the potential to enhance our understanding of both the evolution of animal personality and the ‘winner and loser effects’ of post-fight changes in RHP.
contest; boldness; resource-holding potential; sea anemone; personality; fighting
The outcomes of agonistic interactions modulate access to resources and thereby affect fitness. Success in agonistic encounters may depend on intrinsic physical and physiological performance, and on social experience. Here we test the hypothesis that previous experience will override physical strength in determining the outcome of fights in the freshwater crayfish Cherax dispar. Between unfamiliar opponents, greater chelae closing force significantly increases the chances of winning. However, even when the chelae of the original winners were disabled, the winners kept on winning against the same opponents after 30 min and 24 h. This winner effect disappeared when previous winners encountered unfamiliar individuals. Similarly, a previous loss did not affect the outcomes of subsequent encounters with unknown crayfish. We suggest that this prolonged recognition of individuals and their relative fighting ability is a mechanism that can reduce the number of agonistic encounters experienced by individuals.
aggressive behaviour; fitness; winner effect; social dominance; crayfish
Aggression is ubiquitous in the animal kingdom, whenever the interests of individuals conflict. In contests between animals, the larger opponent is often victorious. However, counter intuitively, an individual that has little chance of winning (generally smaller individuals) sometimes initiates contests. A number of hypotheses have been put forward to explain this behaviour, including the ‘desperado effect’ according to which, the likely losers initiate aggression due to lack of alternative options. An alternative explanation suggested recently is that likely losers attack due to an error in perception: they mistakenly perceive their chances of winning as being greater than they are. We show that explaining the apparently maladaptive aggression initiated by the likely loser can be explained on purely economic grounds, without requiring either the desperado effect or perception errors. Using a game-theoretical model, we show that if smaller individuals can accurately assess their chance of winning, if this chance is less than, but close to, a half, and if resources are scarce (or the contested resource is of relatively low value), they are predicted to be as aggressive as their larger opponents. In addition, when resources are abundant, and small individuals have some chance of winning, they may be more aggressive than their larger opponents, as it may benefit larger individuals to avoid the costs of fighting and seek alternative uncontested resources.
fighting; aggression; Napoleon complex; game theory
Theoreticians predict that animal ‘personality’ traits may be maladaptive if fixed throughout different contexts, so the present study aimed to test whether these traits are fixed or plastic. Rainbow trout (Onchorhyncus mykiss) were given emboldening or negative experiences in the forms of watching bold or shy individuals responding to novelty or winning or losing fights to examine whether prior experience affected boldness. Bold individuals that lost fights or watched shy demonstrators became more shy by increasing their latency to approach a novel object, whereas shy observers that watched bold demonstrators remained cautious and did not modify their responses to novelty. Shy winners became bolder and decreased their latency to approach a novel object, but shy losers also displayed this shift. In comparison, control groups showed no change in behaviour. Bold fishes given negative experiences reduced their boldness which may be an adaptive response; however, shy fishes may base their strategic decisions upon self-assessment of their relative competitive ability and increase their boldness in situations where getting to resources more quickly ensures they outcompete better competitors.
individuality; social status; novelty; prior experience; Salmonidae
Winner and loser effects are defined as an increased probability of winning an aggressive interaction at time T, based on victories at time T-1, T-2, etc., and an increased probability of losing at time T, based on losses at time T-1, T-2, etc., respectively. Prior theoretical work on dominance hierarchy formation has demonstrated that when players are not capable of individual recognition, loser effects always produce a clear top-ranked (alpha) individual, but all other ranks in a group remain unclear; whereas winner effects always produce strict linear hierarchies in which the rank of each individual is clear. Paradoxically, however, when individual recognition--a phenomenon long thought to stabilize hierarchies--is possible, winner and loser effects have no impact on the probability of forming strict linear hierarchies.
Alpha-synuclein (α-Syn) is a small neuronal protein that has been found to be expressed throughout the brain. It has been shown that α-Syn regulates the homeostasis of monoamine neurotransmitters and is involved in various degenerative and affective disorders. There is indication that α-Syn may regulate expression of the brain-derived neurotropic factor (BDNF) which plays an important role in the mood disorders.
The study aimed to analyze the mRNA levels of Snca and Bdnf genes in the ventral tegmental area (VTA) and raphe nuclei of the midbrain in male mice that had each won or defeated 20 encounters (20-time winners and 20-time losers, respectively) in daily agonistic interactions. Groups of animals that had the same winning and losing track record followed by a no-fight period for 14 days (no-fighting winners and no-fighting losers) were also studied. Snca mRNA levels were increased in the raphe nuclei in the 20-time losers and in the VTA of the 20-time winners. After no-fight period Snca mRNA levels decreased in both groups. Snca mRNA levels were similar to the control level in the VTA of the 20-time losers and in the raphe nuclei of the 20-time winners. However Snca gene expression increased in these areas in the no-fighting winners and no-fighting losers in comparison with respective mRNA levels in animals before no-fight period. Bdnf mRNA levels increased in VTA of 20-time winners. Significant positive correlations were found between the mRNA levels of Snca and Bdnf genes in the raphe nuclei.
Social experience affects Snca gene expression depending on brain areas and functional activity of monoaminergic systems in chronically victorious or defeated mice. These findings may be useful for understanding the mechanisms of forming different alpha-synucleinopathies.
A simple three-player model is presented for the evolution of coalitions. The model demonstrates that, under certain conditions, 'winner' and 'loser' effects both favour coalition formation. Winner effects are defined as an increased probability of winning at time T + 1, given a victory at time T, whereas loser effects entail an increased probability of losing at time T + 1, given a loss at time T. Increasing the strength of loser effects or winner effects, or the strength of an individual's position in the hierarchy, makes coalition formation in general more likely, whereas increasing the costs of giving aid does the opposite. The model does not assume any form of reciprocity, but rather examines whether some form of reciprocity or pseudoreciprocity emerges from the model itself. When either winner or loser effects exist, reciprocal coalition formation (e.g. i helps j against k, and j helps i against k) between β (second-ranked individual) and α (highest-ranked individual) or between α and γ (lowest-ranked individual) was possible, but reciprocal aid-giving between γ and β was never favoured. Thus, we have the counterintuitive result that although a coalition between the two lowest members of a hierarchy against the dominant individual is possible (as selection may favour γ aiding β against α), such a coalition is not predicted to be reciprocal in kind. Interpopulational comparisons examining winner/loser effects and coalition formation would allow for a test of many of the model's most basic predictions. Unfortunately, most work on coalitions has been undertaken in primates, whereas work on winner and loser effects has focused on rodents, and more recently, in fish and birds. Hopefully, the model presented here will spur future work that will look at all of these factors simultaneously in many taxa.
Assessment strategies are an important component in game theoretical models of contests. Strategies can be either based on one’s own abilities (self assessment) or on the relative abilities of two opponents (mutual assessment). Using statistical methodology that allows discrimination between assessment types, we examined contests in the jumping spider Phiddipus clarus. In this species, aggressive interactions can be divided into ‘pre-contact’ and ‘contact’ phases. Pre-contact phases consist of bouts of visual and vibratory signaling. Contact phases follow where males physically contact each other (leg fencing). Both weight and vibratory signaling differences predicted winners with heavier and more actively signaling males winning more contests. Vibratory behaviour predicted pre-contact phase duration, with higher signaling rates and larger differences between contestants leading to longer pre-contact interaction times. Contact phase duration was predicted most strongly by the weight of losing males relative to that of winning males, suggesting that P. clarus males use self-assessment in determining contest duration. While a self-assessment strategy was supported, our data suggest a secondary role for mutual assessment (“partial mutual assessment”). After initial contest bouts, male competitors changed their behaviour. Pre-contact and contact phase durations were reduced while vibratory signaling behaviour in winners was unchanged. In addition, only vibratory signaling differences predicted winners in subsequent bouts suggesting a role of experience in determining contest outcomes. We suggest that the rules and assessment strategies males use can change depending on experience and that assessment strategies are likely a continuum between self- and mutual assessment.
The processes that underlie the formation of the dominance hierarchy in a group are since long under debate. Models of self-organisation suggest that dominance hierarchies develop by the self-reinforcing effects of winning and losing fights (the so-called winner-loser effect), but according to ‘the prior attribute hypothesis’, dominance hierarchies develop from pre-existing individual differences, such as in body mass. In the present paper, we investigate the relevance of each of these two theories for the degree of female dominance over males. We investigate this in a correlative study in which we compare female dominance between groups of 22 species throughout the primate order. In our study female dominance may range from 0 (no female dominance) to 1 (complete female dominance). As regards ‘the prior attribute hypothesis’, we expected a negative correlation between female dominance over males and species-specific sexual dimorphism in body mass. However, to our surprise we found none (we use the method of independent contrasts). Instead, we confirm the self-organisation hypothesis: our model based on the winner-loser effect predicts that female dominance over males increases with the percentage of males in the group. We confirm this pattern at several levels in empirical data (among groups of a single species and between species of the same genus and of different ones). Since the winner-loser effect has been shown to work in many taxa including humans, these results may have broad implications.
Winner and loser effects have now been documented in a number of species. To our knowledge, experimental work, however, has focused exclusively on pairwise interactions, and not the extent to which winner and loser effects impact hierarchy formation. We report the results of experimentally manipulated winner and loser effects on hierarchy formation in a socially living species, the green swordtail, Xiphophorus helleri. Our results demonstrate that randomly chosen winners in pairwise aggressive contests were more likely to emerge as top-ranked individuals in a hierarchy, whereas randomly chosen losers were more likely to emerge as the lowest-ranking individuals, and that 'winner-neutral-loser' hierarchies were significantly overrepresented.
Dominance hierarchies pervade animal societies. Within a static social environment, in which group size and composition are unchanged, an individual's hierarchy rank results from intrinsic (e.g. body size) and extrinsic (e.g. previous experiences) factors. Little is known, however, about how dominance relationships are formed and maintained when group size and composition are dynamic. Using a fusion–fission protocol, we fused groups of previously isolated shore crabs (Carcinus maenas) into larger groups, and then restored groups to their original size and composition. Pre-fusion hierarchies formed independently of individuals' sizes, and were maintained within a static group via winner/loser effects. Post-fusion hierarchies differed from pre-fusion ones; losing fights during fusion led to a decline in an individual's rank between pre- and post-fusion conditions, while spending time being aggressive during fusion led to an improvement in rank. In post-fusion tanks, larger individuals achieved better ranks than smaller individuals. In conclusion, dominance hierarchies in crabs represent a complex combination of intrinsic and extrinsic factors, in which experiences from previous groups can carry over to affect current competitive interactions.
aggression; carry-over effects; dominance hierarchy; fission–fusion; social environment
Male mating success is often determined by territory ownership and traits associated with successful territory defense. Empirical studies have shown that the territory owner wins the majority of fights with challenging males. Several physical and physiological traits have been found to correlate with resource holding potential. In addition, in aerial insects, wing design may also have a strong influence on resource holding potential, since it determines efficiency and precision during flight. However, this possibility has not yet been thoroughly evaluated using the modern technique of geometric morphometrics to analyze shape. Therefore, this study examined whether wing shape affects the outcome of male-male contests in the territorial damselfly, Calopteryx virgo (L.) (Odonata: Calopterygidae). Wing shape and also traditional flight-related morphological measures were compared between 27 pairs of winners and losers from experimental territorial contests. Contrary to expectations, there were no differences between winners and losers in all studied wing traits (shape, length, width, total surface, aspect ratio, and wing loading). However, highly significant differences in wing shape and size were detected between the fore- and hindwing. It is currently not known how these differences relate to flight performance, since previous biomechanical studies in damselflies assumed fore- and hindwings to have an identical planform.
aspect ratio; geometric morphometrics; male-male competition; resource holding potential; wing design
Evolutionary processes can interact with the mechanisms of steroid hormone action to drive interspecific variation in behavioural output, yet the exact nature of these interactions is poorly understood. To investigate this issue, we compare the endocrine machinery underlying the winner effect (an ability to increase winning behaviour in response to past victories) in two closely related species of Peromyscus mice. Typically, after winning a fight, California mice (Peromyscus californicus) experience a testosterone (T) surge that helps enhance their future winning behaviour, whereas white-footed mice (Peromyscus leucopus) experience neither a T surge nor a change in subsequent winning behaviour. However, our results indicate that when the post-victory T response of male white-footed mice is phenotypically engineered to resemble that of California mice, individuals are capable of developing a strong and lasting winner effect. Moreover, this ‘induced’ winner effect in white-footed mice qualitatively matches the winner effect that develops naturally in California mice. Taken together, these findings suggest that white-footed mice have the physiological machinery necessary to form a robust winner effect comparable to that formed by California mice, but are unable to endogenously activate this machinery after achieving winning experiences. We speculate that evolutionary processes, like selection, operate on the physiological substrates that govern post-victory T release to guide divergence in the winner effect between these two species.
social behaviour; winner effect; testosterone; behavioural evolution; the challenge hypothesis; Peromyscus mice
Animals are capable of using information from recent experiences to modify subsequent behavioral responses. Animals' ability or propensity to modify their behavior in the light of new information has repeatedly been shown to correlate with, or be influenced by, either their intrinsic competitive ability or their dominance experience - an influence which can be long-lasting. Using a mangrove killifish, Kryptolebias marmoratus, as the study organism, we investigated whether and if so how the effect of a winning or a losing experience one day prior to a dyadic contest was modulated by both competitive ability measured two months previously and a winning or losing experience forced on the contestants one month previously.
Winning/losing experience forced on the fish one month previously affected how they utilized information from their winning/losing experience one day before Test Day: Individuals that were randomly assigned a losing experience one month previously were more susceptible to the influence of their 1-day winning/losing experience than those assigned a winning experience. Competitive ability measured two months previously, winning/losing experience from one month previously and the winning/losing experience received one day previously all significantly influenced the fish's contest behaviors on Test Day, although only 2-month competitive ability significantly influenced escalation duration, indicating that it was still a good index for the fish's competitive ability two months later.
These results suggest that the value to the fish of information from a recent win or loss depends on the outcome of their past contests and show that contest experience has a long-term effect on contest behavior.
animal contest; winner-loser effect; information; fighting ability; Kryptolebias marmoratus
Syrian hamsters readily form dominant-subordinate relationships under laboratory conditions. Winning or losing in agonistic encounters can have striking, long-term effects on social behavior, but the mechanisms underlying this experience-induced behavioral plasticity are unclear. The present study tested the hypothesis that changes in brain-derived neurotrophic factor (BDNF) may at least in part mediate this plasticity. Male hamsters were paired for 15-min using a resident-intruder model, and individuals were identified as winners or losers on the basis of their behavior. BDNF was examined with in situ hybridization 2 hours after treatment during the consolidation period of emotional learning. Losing animals had significantly more BDNF mRNA in the basolateral (BLA) and medial (MeA) nuclei of the amygdala when compared to winning animals as well as novel cage and home cage controls. Interestingly, winning animals had significantly more BDNF mRNA in the dentate gyrus of the dorsal hippocampus (DHPC DG) than did losing animals, novel and home cage controls. No conflict-related changes in BDNF mRNA were observed in several other regions including the bed nucleus of the stria terminalis and central amygdala. Next, we demonstrated that K252a, a Trk receptor antagonist, significantly reduced the acquisition of conditioned defeat when administered within the BLA. These data support a model in which BDNF-mediated plasticity within the BLA supports learning of submission or subordinate social status in losing animals, whereas BDNF-mediated plasticity within the hippocampus may instantiate aspects of winning such as control of a territory in dominant animals.
stress; plasticity; basolateral amygdala; hippocampus; submission
Game theory models of animal contests make many non-mutually exclusive predictions, complicating empirical tests. These predictions regard the relationship between contest parameters and fighting ability, for which body size is usually used as a proxy. However, in many systems, body size may be a limited proxy since multiple traits and contextual factors such as experience influence fighting ability. Using contests between male Cape dwarf chameleons, Bradypodion pumilum, I test alternative game theory models of extended contests. I show how the most likely candidate model can be identified through a process of elimination, based on tests of key predictions. In addition, I present a measure of fighting ability based on multiple traits that allows ability to change as experience changes. In dwarf chameleons, persistence is based on loser thresholds rather than assessment of relative ability, ruling out the sequential assessment model. Winners and losers do not match behaviours in early parts of the contest, arguing against all types of war of attrition models. Although the cumulative assessment model remained as the most likely candidate model, not all specific predictions of this model were upheld.
cumulative assessment; sequential assessment; war of attrition; resource-holding potential; contest competition; signalling
We examined the impact of winner and loser effects on dominance hierarchy formation when individuals are capable of estimating their opponent's resource holding power (RHP). The accuracy of such estimates was a variable in our simulations, and we considered cases in which all individuals err within the same bounds, as well as cases in which some individuals consistently overestimate, while others consistently underestimate their opponent's fighting RHP. In all cases, we found a clearly defined linear hierarchy. In most simulations, the vast majority of interactions were ‘attack–retreats’, and the remainder of interactions were almost all ‘fights’. Error rates had no effect on the linearity of the hierarchy or the basic attack–retreat nature of interactions, and consistent over and underestimation did not affect the ultimate position of an individual in a hierarchy.
winner effect; loser effect; individual recognition
BACKGROUND: Despite the wealth of evidence linking low income to ill health, there is little information from randomised studies on how much and how quickly these risks can be reversed by improvements in income. OBJECTIVE: To conduct a systematic review of randomised studies of income supplementation, with particular reference to health outcomes. DESIGN: Extensive searches of electronic databases and contact with previous authors. As well as searching for trials that were specifically designed to assess the effects of increased income, studies of winners and losers of lotteries were also sought: if winning is purely chance, such studies are, in effect, randomised trials of increased income. RESULTS: Ten relevant studies were identified, all conducted in North America, mostly in the late 1960s and 1970s. Five trials were designed to assess the effects of income supplementation on workforce participation and randomised a total of 10,000 families to 3- 5 years of various combinations of minimum income guarantees and reduced tax rates. Two trials were designed to assess re-offending rates in recently released prisoners and randomised a total of 2400 people to 3-6 months of benefits. One trial was designed to assess housing allowances and randomised 3500 families to three years of income supplements. One trial assessed the health effects of 12 months of income supplementation in 54 people with severe mental illness. Finally, one study compared three groups of people who won different amounts of money in a state lottery. In all these studies the interventions resulted in increases in income of at least one fifth. However, no reliable analyses of health outcome data are available. CONCLUSIONS: Extensive opportunities to reliably assess the effects of increases in income on health outcomes have been missed. Such evidence might have increased the consideration of potential health effects during deliberations about policies that have major implications for income, such as taxation rates, benefit policies, and minimum wage levels. Randomised evidence could still be obtained with innovative new studies, such as trials of full benefit uptake or prospective studies of lottery winners in which different sized winnings are paid in monthly installments over many years.