Winning fights elevates testosterone levels in California mice and enhances future ability to win fights

The 'winner effect' has been studied in a variety of species, but only rarely in mammals. We compared effects of winning three, two, one, or zero resident-intruder encounters on the likelihood of winning a subsequent aggressive encounter in the California mouse (Peromyscus californicus). During the training phase, we ensured that resident males won all encounters by staging contests with mildly sedated, smaller intruders. During the test phase, the resident male encountered an unfamiliar, more evenly matched intruder that had experience winning an encounter and was larger than the resident. Testosterone (T) plasma levels significantly increased after the final test when they had experienced two prior winning encounters, and the probability of winning a future encounter increased significantly after three prior wins independent of intrinsic fighting ability. We hypothesize a 'winner-challenge' effect in which increased T levels serve to reinforce the winner effect in male California mice.     

Effects of Warming Temperatures on Winning Times in the Boston Marathon

It is not known whether global warming will affect winning times in endurance events, and counterbalance improvements in race performances that have occurred over the past century. We examined a time series (1933–2004) from the Boston Marathon to test for an effect of warming on winning times by men and women. We found that warmer temperatures and headwinds on the day of the race slow winning times. However, 1.6°C warming in annual temperatures in Boston between 1933 and 2004 did not consistently slow winning times because of high variability in temperatures on race day. Starting times for the race changed to earlier in the day beginning in 2006, making it difficult to anticipate effects of future warming on winning times. However, our models indicate that if race starting times had not changed and average race day temperatures had warmed by 0.058°C/yr, a high-end estimate, we would have had a 95% chance of detecting a consistent slowing of winning marathon times by 2100. If average race day temperatures had warmed by 0.028°C/yr, a mid-range estimate, we would have had a 64% chance of detecting a consistent slowing of winning times by 2100.     

Persistence of Winner and Loser Effects Depends on the Behaviour Measured

Recent contest experience can influence an individual’s behaviour in subsequent contests. When the probability of winning a subsequent contest is used to quantify experience effects, a loser effect usually lasts longer than a winner effect. This conclusion, however, may be caused by this probability understating the persistence of the influence of a winning experience on contest decisions. Using Kryptolebias marmoratus, a mangrove killifish, as the study organism, we investigated whether different conclusions about the relative persistence of winning and losing experiences would be reached when different aspects of contest behaviour (probability of initiating attacks, probability of winning non‐escalated and escalated contests, escalation rate and contest duration) were measured. The results indicated that the apparent persistence of the effect of winning or losing experiences varied with the behaviour studied. When the likelihood to initiate attacks was used, no winner effect was detected while the loser effect lasted for <1 d. When escalation rate was used, the winner effect lasted for 2–4 d, while the loser effect lasted for 1–2 d. When the probability of winning non‐escalated contests was used, the winner effect was detectable for <1 d, while the loser effect lasted for 2–4 d. And, when contest duration was used, the winner effect was detectable for 2–4 d, but no loser effect was detectable. These results show that (1) the probability of winning a subsequent contest understated the persistence of the influence of a winning experience on the fish’s contest decisions, (2) the measures most effective at detecting winner effects are different from those most effective at detecting loser effects and (3) in K. marmoratus, both effects can be detected 2 d after the completion of experience training but both dissipate in 4 d.     

No effect of blue on winning contests in judo

A study by Rowe et al. reported a winning bias for judo athletes wearing a blue outfit relative to those wearing a white one during the 2004 Olympics. It was suggested that blue is associated with a higher likelihood of winning through differential effects of colour on opponent visibility and/or an intimidating effect on the opponent. However, we argue that there is no colour effect on winning in judo. We show that alternative factors, namely allocation biases, asymmetries in prior experience and differences in recovery time are possible confounding factors in the analysis of Rowe et al. After controlling for these factors, we found no difference in blue and white wins. We further analysed contest outcomes of 71 other major judo tournaments and also found no winning bias. Our findings have implications for sports policy makers: they suggest that a white-blue outfit pairing ensures an equal level of play.     

Can Individual Differences in History of Dominance Explain the Development of Linear Dominance Hierarchies?

Observed dominance hierarchies are often more linear than expected from randomly-formed dominance relationships, and in triads of animals attacks are distributed non-randomly. I hypothesize that an individual's history of dominance affects its probability of initiating aggressive interactions in the future and that individuals with winning records are more likely to initiate (winning begets initiating). Consistent with this hypothesis, evidence is presented that dominant individuals are more likely to attack than subordinate individuals. The winning begets initiating hypothesis may also explain why correlations between predicted dominance ranks (based on size, age etc.) and observed dominance ranks can be low: If the cost of engaging in and losing an interaction is high relative to the potential benefits of winning, then a large individual conditioned to be subordinate may refrain from contesting smaller, dominant individuals despite its actual competitive superiority.     

The ‘home advantage’ is necessary for a full winner effect and changes in post-encounter testosterone

Winning aggressive contests can both enhance future winning ability and change post-encounter hormones; however, it remains unclear if the context of a fight also influences such winner effects and hormone changes. We investigated this issue by using California mice (Peromyscus californicus) to test if the effect of residency status is necessary to improve future winning ability and alter post-encounter hormones. Male mice were subjected to an aggressive contest and their blood was collected 45 min after the fight. Upon contest initiation, focal mice had a ‘home advantage’ and three prior winning experiences, only one of these factors, or neither factor. Only individuals with a ‘home advantage’ and prior winning experience showed a full winner effect. Post-encounter changes in testosterone and progesterone resulted from an interaction between residency status and winning experience. These data indicate that in male California mice a ‘home advantage’ is necessary to produce the full winner effect and that residency status helps regulate inter-individual variation in the expression of post-encounter testosterone pulses and progesterone changes. Furthermore, these respective behavioral and physiological phenomena might be modulated in a context-specific manner, in part by the surrounding physical environment.     

Species differences in the winner effect disappear in response to post-victory testosterone manipulations

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.     

Human group behavior: the ideal free distribution in a three-patch situation

A group of 15 college students was exposed to repeated trials of a task in which money was available for choosing among three colors (blue, red, and green). The amount of winning tokens for each color was varied across phases to test whether group distribution would track the ratio of winning tokens between patches. Confirming previous reports on ideal free performance in humans, group choice proved sensitive to the available resources but tended to undermatch the ratio of winning tokens. The difference-equalization rule of Sokolowski, Tonneau, and Freixa i Baqué [Psychonom. Bull. Rev. 6 (1999) 157] gave a satisfactory fit to the data.     

Previous Experiences Alter the Outcome of Aggressive Interactions Between Males in the Crayfish, Procambarus Clarkii

Various measures of size have been associated with increased likelihood of winning such agonistic encounters between decapod crustaceans. Recent social experience, in the form of 'winner' or 'loser' effects, has also been shown to alter the outcome of encounters between size-matched individuals. This study further explores the importance of social experience in crustacean agonistic encounters. Crayfish, Procambarus clarkii , of different size classes were given a series of winning or losing experiences to determine whether this would affect their likelihood of success in subsequent fights. Recent wins or losses in a series of fights did alter the likelihood of an individual winning a subsequent encounter, despite an apparent disparity in size between opponents that would otherwise serve as an accurate predictor of the outcome.     

The winner and loser effect: integrating multiple experiences

An important question in state-dependent behaviour is how multiple influences on state are integrated to determine current behaviour. Aggressive behaviour is known to be affected by a prior contest experience. Nevertheless, whether and how multiple prior fighting experiences are integrated into a fighting decision remain unexplored. In this study, individuals of Rivulus marmoratus (Cyprinodontidae), a hermaphroditic fish, were given different combinations of two prior fighting experiences to investigate: (1) the effect of penultimate experiences on the probability of winning a subsequent contest; (2) the relative effect of a recent win and loss; and (3) whether the effect of a winning experience was as short lived as observed in other species. Penultimate and recent fighting experiences were given to the test fish approximately 48 and 24 h prior to the dyadic contests, respectively. From the results of the five types of contests staged, we conclude that: (1) penultimate fighting experiences had a significant effect on the probability of winning a subsequent contest; (2) a more recent experience had a more pronounced effect than an earlier experience, which suggested that the effect of a fighting experience would decay and/or the effect of a recent experience would interfere with the effect of an earlier experience; (3) no asymmetric effect between a winning experience and a losing experience was detected; and (4) the effect of both a winning and a losing experience lasted for at least 48 h in R. marmoratus which was the maximum time tested in these experiments. The possible reasons for the differences in results among studies of experience effects on contest outcomes are discussed. Copyright 1999 The Association for the Study of Animal Behaviour.     

Winning and losing: causes for variability in outcome of fights in male Magellanic penguins (Spheniscus magellanicus)

Game theory models predict that fighting ability should be moreimportant in contest outcome when the payoffs of winning arehigh for both contestants, and ownership should be more importantwhen payoffs are low. Male Magellanic penguins (Spheniscusmagellanicus) provide an opportunity to test these predictionsin a natural setting because payoffs of winning are higher for penguins fighting before egg laying and lower for penguinsfighting after egg laying, allowing the prediction of differencesin who should win and lose. We watched an area of approximately2000 Magellanic penguin nests from 1992 to 1996 at Punta Tombobreeding colony, Argentina; we quantified fighting behavior,banded contestants, measured their body size (here used as anindex of fighting ability), determined ownership status whenpossible, and monitored their reproductive success. We determinedthat male Magellanic penguins fought for nests and mates. Astheory predicts, before egg laying, body size difference wasmore important than ownership as a predictor of contest outcome and fight duration. After egg laying, owners won fights, andsize did not predict who won or how long they fought. Our comparisonsof nest ownership, nest quality, and chicks fledged by winnersand losers suggested that our predictions on the change inbenefits of winning before and after egg laying were correct.We conclude that game theory models are useful in predictingwho won or lost fights in male Magellanic penguins and thatultimate benefits of winning fights are related to fitness.     

Why are small males aggressive?

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.     

Imperfect assessment and limited information preclude optimal strategies in male-male fights in the orb-weaving spider Metellina mengei

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.     

An effect of inter-trial duration on the gambler's fallacy choice bias

The gambler's fallacy is defined as the avoidance of a winning outcome in a stochastic environment with a constant probability. We tested the possibility that the gambler's fallacy in humans is responsive to the amount of time between choice allocations. Two groups of subjects were placed in a six-choice betting game in which the choices were clustered into two “patches.” Groups were defined by the length of time - 2 s or 6 s - between trials. On any given trial subjects allocated six points among the alternatives, and retained any points that were bet on the winning alternative. Both groups showed evidence of the gambler's fallacy bias. However, the bias was stronger in the 6-s ITI group than in the 2-s ITI group. This difference was found primarily to be due to differences in the number of subjects showing an opposing bias to the gambler's fallacy, namely a preference for the most recent winning alternative. This choice bias is termed the hot hand fallacy. Our findings contradict predictions derived from a foraging heuristic and from traditional accounts of the gambler's fallacy.     

Cooperation Improves Success during Intergroup Competition: An Analysis Using Data from Professional Soccer Tournaments

The benefit mutually gained by cooperators is considered the ultimate explanation for why cooperation evolved among non-relatives. During intergroup competition, cooperative behaviours within groups that provide a competitive edge over their opposition should be favoured by selection, particularly in lethal human warfare. Aside from forming larger groups, three other ways that individuals within a group can cooperate to improve their chances of gaining a mutual benefit are: (i) greater networking, (ii) contributing more effort, and (iii) dividing labour. Greater cooperation is expected to increase the chances of gaining a group benefit by improving proficiency in the tasks critical to success—yet empirical tests of this prediction using real-world cases are absent. In this study, we used data derived from 12 international and professional soccer competitions to test the predictions that: 1) greater levels of cooperative behaviour are associated with winning group contests, 2) the three forms of cooperation differ in relative importance for winning matches, 3) competition and tournament-type affect the levels of cooperation and shooting proficiency in matches, and 4) greater levels of networking behaviour are associated with increased proficiency in the most critical task linked with winning success in soccer—shooting at goal. Winners were best predicted by higher shooting proficiency, followed by greater frequencies of networking interactions within a team but unexpectedly, fewer networking partners and less division of labour. Although significant variation was detected across competitions and tournament-types, greater levels of networking behaviour were consistently associated with increased proficiency in shooting at goal, which in turn was linked with winning success. This study empirically supports the idea that intergroup competition can favour cooperation among non-relatives.     

Winner and loser effects in human competitions. Evidence from equally matched tennis players

Animals winning an agonistic encounter are more likely to win their next encounter while losers are less likely, even when controlling for motivation and physical size. Do these winner and loser effects exist in human competitions? Drawing on a large database of professional tennis matches, we were able to control for players' ability and thereby test for winner and loser effects. We narrowed the database to matches between players who on average did not differ significantly in rank, and further to matches in which the first set was fought to a long tie-break. These closely fought matches present a natural experiment because players are assigned to treatment conditions – winning or losing a set – despite similar ability and performance. We found that among men, the winner of a closely fought tie-break had an approximate 60% chance of winning the second set, the loser a 40% chance. These effects did not exist among women, a finding consistent with the hypothesis that androgens mediate winner and loser effects. Our results may help in the design of competitions in sport as well as in work environments, where it may prove useful to either encourage winner effects or to attenuate their occasional adverse consequences.     

Changes in Heart Rate Associated with Contest Outcome in Agonistic Encounters in Lobsters

Agonistic contests between lobsters housed together in a confined space progress through encounters of increasing intensity until a dominance relationship is established. Once this relationship is established, losing animals continually retreat from the advances of winners.These encounters are likely to consume much energy in both winning and losing animals. Therefore, one might expect involvement of many physiological systems before, during and after fights. Here, we report effects of agonistic encounters on cardiac frequency in winning and losing adult lobsters involved in dyadic interactions.The results show that: (i) small but significant increases in heart rate are observed upon chemical detection of a conspecific; (ii) during agonistic interactions, further increases in heart rate are seen; and (iii) ultimate winners exhibit greater increases in heart rate lasting longer periods of time compared to ultimate losers. Heart rate in winners remains elevated for at least 15 min after the contests have ended and animals have been returned to their home tanks. Reduced effects are seen in second and third pairings between familiar opponents.The sustained changes in heart rate that we observe in winning lobsters may result from hormonal modulation of cardiac function related to the change in social status brought about by contest outcome.     

Fighting and aggressive sound determines larger male to win male-male competition in a bark beetle

Intrasexual selection occurs in male-male competition over access to females and usually results in the larger male winning.While much research has documented that size matters,little is known about how the larger male wins.Dendroctonus vale ns is an aggregating monogamous bark beetle in which males have large variation in body size and display intense competition over females.Behavioral observation showed two males fight each other within the gallery by pushing/shoving and stridulated more when two males encountered each other.Experiments using two different-sized males synchronously competing showed that larger males won 95%of contests.Reciprocal displacement experiments using muted and intact males of different or equal size were used to simulate male-male competition.Larger males displaced the smaller resident male in 90%of contests,while smaller males prevailed over larger residents in 6.7%of contests.With both males silenced,larger males displaced smaller males in 80%of con tests,while smaller males prevailed in 8%of con tests.Further experiments using equal-sized males showed aggressive soundemitting males displaced muted males in 67%of contests,yet intact males displaced other intact males in only 37.5%of con tests.Sound analysis showed sound pressure level is an honest signal of body size and males chose soft sounds over loud aggressive sounds in assays.Therefore,D.valens males have evolved dual behaviors,fighting and aggressive sounds associated with body size,to assess rivals to compete for a partner,gaining insights in male-male competition for this species and for other animals.     

Experimental evidence that winning or losing a fight does not affect sperm quality in a field cricket

Fighting is a powerful social experience that can affect male reproductive behavior, including ejaculatory strategies. Whereas winners may monopolize females, losers may instead perceive high sperm competition and limited future mating opportunities, and accordingly enhance ejaculate quality to maximize their reproductive success. In male field crickets Gryllus bimaculatus that fight aggressively for control of breeding territories, winners are known to possess sperm of lower quality (viability) compared to losers, but it remains unclear whether this is due to short‐term fighting consequences. To test if the fighting experience per se (winning or losing) affects male adjustment of sperm viability, we subjected males to winning and losing experiences by staging fights against size‐matched rivals of known fighting ability. These rivals were males that previously won or lost a fight and, due to “winner‐loser effects” kept winning or losing subsequent contests. We sampled sperm prior and after the fight and twice in control males with no fighting experience and found no differences in sperm viability across measures. We conclude that males do not tailor their ejaculate quality following a single fight, or based on its outcome. Intrinsic differences in other attributes between winners and loser phenotypes may explain differences in sperm quality previously described in this system.     

Winning, losing, mood, and testosterone.

In two experiments, male college students either won or lost $5 on a task controlled entirely by chance. In both studies, winners reported a more positive mood change than did losers and, in Experiment 2, winners reported a more positive mood change than a neutral group that did not win or lose money. After the task was completed, winners exhibited significantly higher testosterone levels than losers. Levels of cortisol, a hormone associated with stress and arousal, did not differ among the groups, suggesting that a hormone-behavior response pattern for winning and losing is specific to testosterone. These data suggest that winning can alter testosterone levels in men and that mood may mediate such changes.