Impaired sperm quality,delayed mating but no costs for offspring fitness in crickets winning a fight

The outcome of male–male contest competition is known to affect male mating success and is believed to confer fitness benefits to females through preference for dominant males. However, by mating with contest winners, females can incur significant costs spanning from decreased fecundity to negative effects on offspring. Hence, identifying costs and benefits of male dominance on female fitness is crucial to unravel the potential for a conflict of interests between the sexes. Here, we investigated males' pre‐ and post‐copulatory reproductive investment and its effect on female fitness after a single contest a using the field cricket Gryllus bimaculatus. We allowed males to fight and immediately measured their mating behaviour, sperm quality and offspring viability. We found that males experiencing a fight, independently of the outcome, delayed matings, but their courtship effort was not affected. However, winners produced sperm of lower quality (viability) compared to losers and to males that did not experience fighting. Results suggest a trade‐off in resource allocation between pre‐ and post‐mating episodes of sexual selection. Despite lower ejaculate quality, we found no fitness costs (fecundity and viability of offspring) for females mated to winners. Overall, our findings highlight the importance of considering fighting ability when assessing male reproductive success, as winners may be impaired in their competitiveness at a post‐mating level.     

Why do winners keep winning? Androgen mediation of winner but not loser effects in cichlid fish

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.     

Ejaculatory strategies associated with experience of losing

Modulation of behaviours as a result of fighting experience has been observed in many animals and can influence pre-copulatory sexual selection. This study investigated how fighting experience affects ejaculatory strategies. In male flour beetles, Gnatocerus cornutus, experience of losing a fight decreases a male''s aggressiveness for up to 4 days. We found that males losing a fight show increased ejaculatory investment, but there was no ejaculatory modulation owing to winning. However, the increase in ejaculate investment following a loss was no longer observed after 5 days. These results indicate that males adjust their investment in sperm competition according to their experience, and that fighting experience can significantly influence pre- and post-copulatory reproductive tactics.     

Fighting dynamics of male copperheads,Agkistrodon contortrix (Serpentes,Viperidae): Stress-induced inhibition of sexual behavior in losers

Adult male copperheads (Agkistrodon contortrix) fight for priority of access to females during the mating periods in spring and late summer. During fights, one male abruptly quits and retreats, and the other chases in pursuit. One male thus emerges as the winner and the other as the loser. Reversal of this outcome does not occur during the time of observation (30 min) nor in 24 hr postfight trials. In all cases, winners gain priority of access to females. Losers, in contrast, do not pursue females nor gain access to them, even when winners are removed from the arena. In this study, courtship performance of male A. contortrix was studied in the laboratory using subjects with either winning or losing experience from staged fights. All males used first were tested with a single female to determine courtship performance prior to the agonistic trials. From these tests, each male was given a single courtship score of 0 (no courtship) to 3 (most intense courtship). Only males receiving a score of 2–3 were used in the initial agonistic trials. All staged fights were conducted in a large arena and involved two males and one female. Following fights, winners and losers were tested again for courtship performance. In trials conducted at 24 hr and 7 days postfight, only losers were tested. It was found that prefight courtship scores were not significantly different between winners and losers. At 30 min postfight, most losers showed complete suppression of courtship behavior (score 0). Winners, in contrast, showed equivalent or an increase in their courtship scores. At 24 hr postfight, courtship scores of losers remained significantly lower than their prefight scores. At 7 days postfight, courtship scores of losers were not significantly difference from their original prefight scores. Fighting behavior in free-ranging A. contortrix represents a potentially significant cost to losers if it is associated with loss of reproductive opportunities. Recent evidence from hormonal studies indicates that inhibition of courtship and fighting behavior in male A. contortrix is stress-induced. © 1996 Wiley-Liss, Inc.     

Influence of previous residency and body mass in the territorial contests of the butterfly Hermeuptychia fallax (Lepidoptera: Satyrinae)

Although many studies have evaluated the possible rules used by males to settle territorial contests, few have simultaneously considered the possibility of mutual or self-assessment of fighting ability and the effect of prior residence on the chances of winning. Here, we experimentally investigated the influence of previous residency on the development and outcome of contests between males of the butterfly Hermeuptychia fallax. By comparing the frequency of resident victories in nonmanipulated disputes versus interactions in which both rivals were induced to behave as residents, we showed that prior residency may interact with body mass in contest resolution. In nonmanipulated disputes, resident males showed high winning probabilities, although they were similar to losers in body mass. However, when both males behaved as residents, the disputes were longer and were settled in favor of the heavier male. This indicates that, although residency is an important component of male willingness to fight, it is not the only factor influencing conflict resolution. Despite the correlation between body mass and territorial success, the weight of neither winners nor losers correlated with fighting duration. We conclude that residence asymmetry may sometimes conceal male traits important to contest resolution and that it may obscure the identification of the assessment strategy used in contests.     

Repeated Recent Aggressive Encounters Do Not Affect Behavioral Consistency in Male Siamese Fighting Fish

Consistent individual differences in behavior suggest that individuals respond in a predictable and repeatable manner in a specific situation while differing from other individuals. Male Siamese fighting fish exhibit consistent individual differences in decision‐making strategies when they encounter a receptive female and a rival male simultaneously. However, whether these differences are altered by recent experience is unknown. We examined the influence of repeated aggressive encounters on behavioral consistency and decision‐making. Males were presented with paired female–male dummies prior to any aggressive experiences to obtain a baseline measure. Next, males either won or lost three consecutive contests against rivals and then received the paired female–male dummies after each of these encounters. Overall levels of highly aggressive behaviors were affected by contest outcome, while levels of female‐directed were not. Not surprisingly, winning a fight led to an increase in male‐directed bites, an overtly aggressive behavior that only occurs after fights have escalated. Fighting a male before encountering the dummies caused males to perform more tail beats to the dummy male, perhaps as a result of increased motivation. Males exhibited similar levels of repeatability and used the same strategies when faced with conflicting stimuli regardless of fighting experience. Thus, while winning or losing a fight impacts overall aggression, it does not influence behavioral consistency. This study demonstrates that consistent individual differences and decision‐making strategies may be resistant to recent aggressive experiences, even over a period of days.     

Assessment Strategies and the Effects of Fighting Experience on Future Contest Performance in the Green Anole (Anolis carolinensis)

Social experiences can be useful sources of information for animals charged with making fitness‐related decisions. Fighting experience can alter an animal's perception of its fighting ability possibly leading to changes in future contest decisions, which may increase/decrease their probability of winning future contests. Winner and loser effects have been revealed in a wide array of animals, but studies using reptilian models are rare. This study investigated the impact of fighting experience on future contest performance and outcome in the green anole lizard and investigated the assessment strategies used by anoles during contests of different intensities. To determine whether the green anole expresses winner or loser effects, focal animals engaged in a primary contest with a smaller (larger) opponent to gain a winning (losing) experience; opponent size asymmetries were a significant predictor of contest outcome. Focal individuals were isolated for 2 d before being given a secondary contest with a size‐matched, naïve opponent. We found no evidence of winner or loser effects 2 d following a previous contest. Although previous contest outcome did not dictate future contest success, dynamics of the previous contest did. Highly aggressive primary contest losers won a significant proportion of the secondary contests, while less aggressive losers were more apt to lose the secondary contest. Secondary contest success of prior winners was not influenced by earlier contest performance. Further analyses of contest dynamics reveal that individuals may use different assessment strategies depending on the intensity of the contest. Our results demonstrate that future contest success may be driven more by individual performance in a prior contest and less by prior contest outcome.     

Rats Benefit from Winner and Loser Effects

Prior fighting experience of opponents can influence the outcome of conflicts. After a victory, animals are more likely to win subsequent contests, whereas after a defeat animals are more likely to lose, regardless of the identity of opponents. The underlying mechanisms and the adaptive significance of these winner and loser effects are as yet unknown. Here, we tested experimentally whether agonistic behavior of male wild‐type Norway rats is influenced by social experience, and we investigated whether this might reduce fighting costs (duration of contest, risk of injury) in subsequent encounters. Rats were randomly assigned to receive either a losing or a winning experience and subsequently tested with unfamiliar, naïve opponents. We found that most rats with a winning experience won the subsequent encounter, and all rats with a losing experience lost the next contest. Previous winners attacked more rapidly in the subsequent encounter and reduced their aggressive behavior sooner; the contests were decided more quickly, which saved time and behavioral effort to the winner. Previous losers received less aggression in the next encounter, despite emitting fewer submissive ultrasonic calls than in the preceding contest, thereby reducing the risk of being injured by the opponent. Thus, anonymous social experience influenced rats’ subsequent behavior toward size‐matched, naïve, unknown social partners. Furthermore, apparently, they benefit from showing winner and loser effects in intraspecific contests by saving time, energy, and risk.     

Male Success in Intrasexual Contests Extends to the Level of Sperm Competition in a Species of Dung Roller Beetle

Contests between males fighting over limited resources often determine the reproductive success of the contestants. It is possible that the outcomes of such contests are affected by traits that also contribute to the likelihood of males achieving paternity when sperm from multiple males compete to fertilize the ova of a single female; however, this relationship has been poorly characterized. In dung and carrion roller beetles, contests among males for the possession of a food ball – a vital resource for nesting – are frequent in nature. However, females that roll and nest with a male often store the sperm of previous mates, forcing males to engage in sperm competition. We analyzed the relationship between wins and losses in contests between food ball owner males and subsequent paternity success in the carrion roller beetle Canthon cyanellus using both naïve (virgin) and experienced (previously mated) males. Owner males that initiated contests were more likely to win, and contest durations were longer between experienced owner males than between naïve owners. The paternity of the winner owner males nesting with a female that had previously mated with another male was higher than for loser owner males, and the paternity of experienced owners also tended to be higher than for virgin owners. Our results indicate that winning and being an experienced male are associated with greater success at fighting and sperm competition than losing and being a naïve male.     

The influence of sex, line, and fight experience on aggressiveness of the Siamese fighting fish in intrasexual competition

We examined the influence of sex, line, i.e., broods from different parents, and previous fight experience on the aggressiveness of the Siamese fighting fish Betta splendens in intrasexual competition. The innate aggressiveness of the fish against their mirror images was measured on the day prior to the direct fight with other individuals, and it was found to be influenced by the line type but not by the sex. In the direct fight with other individuals, the males invested more effort in the fight than the females. In addition, the individuals of a particular line that exhibited a lower innate aggressiveness spent less time in the direct fight and were often losers when compared with those of other lines. After the direct fight with other individuals, the aggressiveness of the fish against their mirror images was remarkably influenced by the outcome of the direct fight, i.e., the winners exhibited more aggressive behavior, whereas the losers exhibited a lesser degree of aggressive behavior. This influence of the previous fight experience on subsequent aggressiveness was the greatest in the individuals of the line that have exhibited the lowest innate aggressiveness. However, the positive effect of the winning experience or the negative effect of the losing experience on subsequent aggressiveness decreased following several days after the previous fight increased.     

Demonstration of Strength and Concealment of Weakness in Escalating Fights of Male Swordtails (Xiphophorus helleri)

In the green swordtail (Xiphophorus helleri) confrontations between strange males regularly escalate to high levels of mutual Bites and Fin Grips, even between males differing greatly in size. The original expectation of early game theory models that the behaviours of the ultimate winners and losers are indistinguishable until shortly before the end of the fight could not be confirmed. A significant characteristic of loser behaviour is that the Biting rates are higher before and after escalation. Conversely, Fin Grips are more frequent in the ultimate winners than the losers. However, such behavioural differences are very poor predictors of the outcome of the fight from the viewpoint of a single fighting individual during the contest. Correct forecasts did not exceed 67%. The escalating fights of swordtails are considered as trials of strength in which the stronger male tries to demonstrate strength by the most costly behaviour pattern available, namely Fin Grips, and the weaker male conceals weakness quite successfully by countering with the same tactic and suppressing signs of weakness such as Avoidance behaviour.     

Does prior residency interact with loss? A study of male–male contests in the hermit crab Pagurus minutus

Residency is an important predictor of success in contests with ownership asymmetries. Residency often can interact with a winning experience. However, given that some residents lose a contest even when showing an ownership advantage and that the process leading to loss often determines the loser's subsequent success, prior ownership might also interact with a loss. Here, we staged experimental contests between males of the hermit crab Pagurus minutus with a similar-sized weapon (i.e., cheliped) to examine this possibility. Male–male contests in this species occur between a solitary intruder and an owner guarding a mature female. We evaluated (a) whether resource ownership and female value affect the contest outcome and (b) whether the probability of winning after losing differs depending on the initial role of the loser (i.e., owner or intruder) by using precopulatory guarding pairs of P. minutus collected from the field. In the first fighting trial, we found an ownership advantage and increasing owner success as the body size of his partner increased. Although some owners lost the fight, in contrast to our prediction, the frequency of losing in the second fighting trial did not differ between prior owners and prior intruders. Because losers from the first fighting trial of male–male contests have no female regardless of their initial role, this shared solitary status might be related to the lack of difference in success in the second fighting trial. Moreover, unlike in other animals, resident status might not always assure greater fighting ability in P. minutus males because guarding Pagurus males can avoid male–male contests by climbing up objects in the field. Losers in the first trial, therefore, may have been weaker contestants based on traits other than size, regardless of whether they were owners or intruders.     

Is boldness a resource-holding potential trait? Fighting prowess and changes in startle response in the sea anemone,Actinia equina

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.     

Examination of prior contest experience and the retention of winner and loser effects

In many animal taxa, prior contest experience affects future performance such that winning increases the chances of winning in the future (winner effect) and losing increases the chances of losing in the future (loser effect). It is, however, not clear whether this pattern typically arises from experience effects on actual or perceived fighting ability (or both). In this study, we looked at winner and loser effects in the jumping spider Phidippus clarus. We assigned winning or losing experience to spiders and tested them against opponents of similar fighting ability in subsequent contests at 1-, 2-, 5-, and 24-h intervals. We examined the strength of winner and loser effects, how long effects persist, as well as how experience affected perceived and actual fighting ability. Our results demonstrate that winner and loser effects are of approximately the same magnitude, although loser effects last longer than winner effects. Our results also demonstrate that previous experience alters actual fighting ability because both the assessment and escalation periods were affected by experience. We suggest that the retention time of experience effects depends on expected encounter rates as well as other behavioral and ecological factors. In systems with short breeding seasons and/or rapidly fluctuating populations, context-dependent retention of experience effects may allow males to track their status relative to the fluctuating fighting ability of local competitors without paying the costs necessary to recall or assess individual competitors.     

Male attractiveness covaries with fighting ability but not with prior fight outcome in house crickets

In many species males that tend to win fights against othermales are more attractive to females. There are three ways inwhich male fighting ability and attractiveness may be associated:(1) attractiveness and fighting ability are influenced by thesame underlying traits (e.g., body size), (2) females prefermales that have directly observed winning fights, or (3) winningprevious fights indirectly improves a male's chance of beingpreferred by females. The last possibility may arise as a consequenceof the "loser effect"; in many species when a male loses a fighthis probability of losing subsequent fights increases. Thereare, however, no studies testing whether such a "loser effect"also influences male attractiveness. Here we show that maleattractiveness and fighting ability are positively correlatedin the house cricket, Acheta domesticus. Our experiment wasdesigned so that females could not directly observe the outcomeof fights, thus eliminating possibility (2) above. We then testedbetween possibilities (1) and (3) by making use of the factthat in some cricket species the "loser effect" can be eliminatedexperimentally by ‘shaking’ a male and stimulatingthe motor program for flying. We showed that in A. domesticus‘shaking’ does affect the outcome of subsequentfights. Males that had won two previous fights were less likelyto win a fight after being ‘shaken’ than when subjectto a control treatment. In contrast, males that had lost twoprevious fights were more likely to win a fight after being‘shaken’ than when they were not shaken. There was,however, no effect of ‘shaking’ on male attractiveness.We conclude that the "loser effect" does not alter the tendencyfor large, dominant males to be attractive to females. Instead,it appears that there are traits correlated with both fightingability and attractiveness. One such trait is body size. Fightwinners were significantly larger than losers and attractivenesswas positively correlated with male body size.     

Effects of physical and social experiences and octopamine receptor agonist on fighting behavior of male crickets Velarifictorus aspersus (Orthoptera: Gryllidae)

Fighting commonly occurs among animals and is very important for resolving conflicts between conspecific individuals over limited resources. The plasticity of fighting strategies and neurobiological mechanisms underlying fighting behavior of insects are not fully understood. In the present study, we examined whether physical and social experiences affected the aggressiveness of males of the cricket Velarifictorus aspersus Walker, and whether an octopamine (OA) receptor agonist could affected the aggressiveness of males exposed to different experiences. We found that flight and winning a fight significantly enhanced male aggressiveness, while losing a fight significantly suppressed male aggressiveness, consistent with the findings of existing studies on other cricket species. We also found that female presence had a stronger enhancing effect on male aggressiveness than flight or winning a fight. These findings demonstrated that physical and social experiences can affect the fighting behavior of male V. aspersus. Topical application of a 0.15?M solution of an OA receptor agonist (chlordimeform, CDM) significantly increased male aggression level, suggesting that OA may play an important role as a neuromodulator in controlling fighting behavior of males of this species. Despite displaying a significantly higher aggression level (level 5 or 6), CDM-treated losers did not escalate to physical combat, while fights between courting males usually resulted in physical escalation. It is likely that fighting behavior is only partly regulated by OA, and additional regulatory pathways may be involved in achieving physical combat.     

Sperm traits negatively covary with size and asymmetry of a secondary sexual trait in a freshwater crayfish

In species where females mate promiscuously, the reproductive success of males depends both on their ability to acquire mates (pre-copulatory sexual selection) and ability of their ejaculates to outcompete those of other males (post-copulatory sexual selection). Sperm competition theory predicts a negative relationship between investment in body traits favouring mate acquisition (secondary sexual characters, SSCs) and investment in ejaculate size or quality, due to the inherent costs of sperm production. In contrast, the phenotype-linked fertility hypothesis posits that male fertilizing efficiency is reliably reflected by the phenotypic expression of male SSCs, allowing females to obtain direct benefits by selecting more ornamented males as copulation partners. In this study, we investigated the relationships between male SSCs and size and quality (viability and longevity) of ejaculates allocated to females in mating trials of the freshwater crayfish Austropotamobius italicus. We showed that the relative size of male weapons, the chelae, was negatively related to ejaculate size, and that chelae asymmetry, resulting from regeneration of lost chelipeds, negatively covaried with sperm longevity. Moreover, males allocated more viable sperm to mates from their own rather than different stream of origin. Our findings thus suggest that, according to sperm competition theory, pre-copulatory sexual selection for large weapons used in male fighting may counteract post-copulatory sperm competition in this crayfish species, and that investment in cheliped regeneration may impair ejaculate quality.     

Seminal fluid affects sperm viability in a cricket

Recent studies have suggested that males may vary the quality of their ejaculates in response to sperm competition, although the mechanisms by which they do so remain unclear. The viability of sperm is an important aspect of ejaculate quality that determines competitive fertilization success in the field cricket Teleogryllus oceanicus. Using in vitro mixtures of sperm and seminal fluid from pairs of male crickets, we show that seminal fluid can affect the viability of sperm in this species. We found that males who invest greatly in the viability of their own sperm can enhance the viability of rival sperm, providing the opportunity for males to exploit the investments in sperm competition made by their rivals. Transitive effects of seminal fluids across the ejaculates of different males are expected to have important implications for the dynamics of male investments in sperm competition.     

Eavesdropping on visual cues in green swordtail (Xiphophorus helleri) fights: a case for networking

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.     

Reciprocity between endocrine state and contest behavior in the killifish, Kryptolebias marmoratus

Given the dramatic behavioral effects of winning and losing contests, and pronounced changes in stress and sex steroid hormones post-fight, it is reasonable to suppose that these hormones also dictate future behavior. We sampled water-borne cortisol, testosterone (T), and 11-ketotestosterone (KT) before and after contests in the mangrove killifish, Kryptolebias marmoratus, to determine how endogenous steroid hormone levels might predict and respond to contest dynamics or success. Pre-fight cortisol related negatively, and pre-fight T related positively to contest initiation and winning, particularly in the smaller opponent. In the pairs where a larger fish won the contest, winners with higher pre-fight T and lower pre-fight cortisol delivered more attacks to the losers. Contest duration and escalation influenced post-fight hormone concentrations primarily in losers. Escalation significantly increased post-fight cortisol, T, and KT for losers but not for winners. However, winners that attacked losers at higher rates had higher levels of post-fight cortisol. Losers also demonstrate the most consistent post-fight hormone responses, particularly to contest escalation and duration. Despite the bidirectional relationship between hormones and contest behavior, we found no overall mean differences in pre- or post-fight cortisol, T, or KT between eventual winners and losers. Thus, it is evident that the categorical states of winner and loser cannot alone reveal the complex, reciprocal associations between endocrine systems and social behavior.