Opponent familiarity and contest experience jointly influence contest decisions in <Emphasis Type="Italic">Kryptolebias marmoratus</Emphasis>

Introduction

Individual recognition and winner/loser effects both play important roles in animal contests, but how their influences are integrated to affect an individual’s contest decisions in combination remains unclear. Individual recognition provides an animal with relatively precise information about its ability to defeat conspecifics that it has fought previously. Winner/loser effects, conversely, rely on sampling information about how an animal’s ability to win compares with those of others in the population. The less precise information causing winner/loser effects should therefore be more useful to an individual facing an unfamiliar opponent. In this study, we used Kryptolebias marmoratus, a hermaphroditic mangrove killifish, to test whether winner/loser effects do depend on opponent familiarity. In addition, as previous studies have shown that subordinates that behave aggressively sometimes suffer post-retreat retaliation from contest winners, we also explored this aspect of contest interaction in K. marmoratus.

Results

In the early stages of a contest, subordinates facing an unfamiliar dominant were more likely to signal their aggressiveness with either gill displays or attacks rather than retreating immediately. A winning experience then increased the likelihood that the most aggressive behavioral pattern the subordinates exhibited would be attacks rather than gill displays, irrespective of their opponents’ familiarity. Dominants that received a losing experience and faced an unfamiliar opponent were less likely than others to launch attacks directly. And subordinates that challenged dominants with more aggressive tactics but still lost received more post-retreat attacks from their dominant opponents.

Conclusions

Subordinates’ contest decisions were influenced by both their contest experience and the familiarity of their opponents, but these influences appeared at different stages of a contest and did not interact significantly with each other. The influence of a losing experience on dominants’ contest decisions, however, did depend on their subordinate opponents’ familiarity. Subordinates and dominants thus appeared to integrate information from the familiarity of their opponents and the outcome of previous contests differently, which warrants further investigation. The higher costs that dominants imposed on subordinates that behaved more aggressively toward them may have been to deter them from either fighting back or challenging them in the future.

     

The Effects of Food Deprivation in Dyadic Encounters in Apodemus sylvaticus

The nature of dominance in relationships among male Apodemus sylvaticus was investigated using multivariate analysis and by experimental manipulation of the behaviour of dominants. The behaviour of dominant mice was typified by greater frequencies and durations of exploration, approach and pursuit of subordinates in short encounters in a small arena. The direction of encounters normally remained constant over 48 h. When the dominant mouse in a dyad was deprived of food there was a change in its behaviour and an immediate change in the behaviour of the subordinate. Subordinates showed greater interest in starving, though known, dominants, explored more freely and indulged more in pursuit than control subordinate mice i. e. those matched with dominants on an adequate diet. Results suggest that in A. sylvaticus, dominance is associated with the inhibition of the activity of a subordinate and the relationship is maintained by the behaviour of the dominant rather than the deference of the subordinate.     

Social conditioning and dominance in male betta splendens

To determine whether social conditioning modifies relative dominance in the Siamese fighting fish ($\text{Betta splendens}$), individual fish were matched for gill cover erection frequency (GCEF) to a mirror. One member of each pair experienced a series of four winning encounters (dominants) while the other of the pair experienced four losing encounters (subordinates). After the fighting experience, matched pairs were placed together and aggressive responses of both fish were recorded. Fish experiencing wins displayed significantly more and were more aggressive than fish experiencing losses which showed significantly higher levels of avoidance. However, no change in mirror GCEF occurred as a result of differential experience.     

Sibling rivalry: training effects, emergence of dominance and incomplete control

Within-brood or -litter dominance provides fitness-related benefits if dominant siblings selfishly skew access to food provided by parents in their favour. Models of facultative siblicide assume that dominants exert complete control over their subordinate sibling's access to food and that control is maintained, irrespective of the subordinate's hunger level. By contrast, a recent functional hypothesis suggests that subordinates should contest access to food when the cost of not doing so is high. Here, we show that within spotted hyena (Crocuta crocuta) twin litters, dominants most effectively skew access to maternal milk in their favour when their aggression prompts a highly submissive response. When hungry, subordinates were less submissive in response to aggression, thereby decreasing lost suckling time and increasing suckling time lost by dominants. In a species where adult females socially dominate adult males, juvenile females were more often dominant than males in mixed-sex litters, and subordinate sisters used more effective counter-tactics against dominant brothers than subordinate brothers against dominant sisters. Our results provide, to our knowledge, the first evidence in a mammal that dominant offspring in twin litters do not exert complete control over their sibling's access to resources (milk), and that sibling dominance relationships are influenced by sibling sex and training effects.     

A Trouble Shared Is a Trouble Halved: Social Context and Status Affect Pain in Mouse Dyads

In mice behavioral response to pain is modulated by social status. Recently, social context also has been shown to affect pain sensitivity. In our study, we aimed to investigate the effects of interaction between status and social context in dyads of outbred CD-1 male mice in which the dominance/submission relationship was stable. Mice were assessed for pain response in a formalin (1% concentration) test either alone (individually tested-IT), or in pairs of dominant and subordinate mice. In the latter condition, they could be either both injected (BI) or only one injected (OI) with formalin. We observed a remarkable influence of social context on behavioral response to painful stimuli regardless of the social status of the mice. In the absence of differences between OI and IT conditions, BI mice exhibited half as much Paw-licking behavior than OI group. As expected, subordinates were hypoalgesic in response to the early phase of the formalin effects compared to dominants. Clear cut-differences in coping strategies of dominants and subordinates appeared. The former were more active, whereas the latter were more passive. Finally, analysis of behavior of the non-injected subjects (the observers) in the OI dyads revealed that dominant observers were more often involved in Self-grooming behavior upon observation of their subordinate partner in pain. This was not the case for subordinate mice observing the pain response of their dominant partner. In contrast, subordinate observers Stared at the dominant significantly more frequently compared to observer dominants in other dyads. The observation of a cagemate in pain significantly affected the observer''s behavior. Additionally, the quality of observer''s response was also modulated by the dominance/submission relationship.     

Better safe than sorry--socio-spatial group structure emerges from individual variation in fleeing, avoidance or velocity in an agent-based model

In group-living animals, such as primates, the average spatial group structure often reflects the dominance hierarchy, with central dominants and peripheral subordinates. This central-peripheral group structure can arise by self-organization as a result of subordinates fleeing from dominants after losing a fight. However, in real primates, subordinates often avoid interactions with potentially aggressive group members, thereby preventing aggression and subsequent fleeing. Using agent-based modeling, we investigated which spatial and encounter structures emerge when subordinates also avoid known potential aggressors at a distance as compared with the model which only included fleeing after losing a fight (fleeing model). A central-peripheral group structure emerged in most conditions. When avoidance was employed at small or intermediate distances, centrality of dominants emerged similar to the fleeing model, but in a more pronounced way. This result was also found when fleeing after a fight was made independent of dominance rank, i.e. occurred randomly. Employing avoidance at larger distances yielded more spread out groups. This provides a possible explanation of larger group spread in more aggressive species. With avoidance at very large distances, spatially and socially distinct subgroups emerged. We also investigated how encounters were distributed amongst group members. In the fleeing model all individuals encountered all group members equally often, whereas in the avoidance model encounters occurred mostly among similar-ranking individuals. Finally, we also identified a very general and simple mechanism causing a central-peripheral group structure: when individuals merely differed in velocity, faster individuals automatically ended up at the periphery. In summary, a central-peripheral group pattern can easily emerge from individual variation in different movement properties in general, such as fleeing, avoidance or velocity. Moreover, avoidance behavior also affects the encounter structure and can lead to subgroup formation.     

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.     

Effect of the Presence of Subordinates on Dominant Female Behaviour and Fitness in Hierarchies of the Dwarf Angelfish Centropyge bicolor

In many hierarchical animal societies, dominant individuals control group membership owing to their power to evict subordinates. In such groups, the presence of subordinates, and therefore group stability, is continually dependent on subordinates being tolerated by dominants. The dominant decision to tolerate or evict is, in turn, dependent on the costs and benefits to dominants of subordinate presence. We investigated the effect of subordinate presence on dominants in the female dominance hierarchy of the dwarf angelfish Centropyge bicolor, using both observations of natural groups and experimental removals of subordinates. We found that the presence of subordinates had no effect on dominant access to resources, as measured by dominant foraging rates and home range areas, nor on dominant fitness, as measured by growth rates and spawning frequencies. Our results suggest that the presence of subordinates has a neutral effect on the current fitness of dominants, so that dominants have no great incentive to evict subordinates. We discuss the possibility that tolerance of subordinates might be further explained by considering future fitness, as dominant females in these haremic protogynous angelfish stand to inherit the male position, whereupon subordinate females change from potential competition to useful mates.     

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.     

Winning agonistic encounters increases testosterone and androgen receptor expression in Syrian hamsters

Winning aggressive disputes is one of several experiences that can alter responses to future stressful events. We have previously tested dominant and subordinate male Syrian hamsters in a conditioned defeat model and found that dominant individuals show less change in behavior following social defeat stress compared to subordinates and controls, indicating a reduced conditioned defeat response. Resistance to the effects of social defeat in dominants is experience-dependent and requires the maintenance of dominance relationships for 14 days. For this study we investigated whether winning aggressive interactions increases plasma testosterone and whether repeatedly winning increases androgen receptor expression. First, male hamsters were paired in daily 10-min aggressive encounters and blood samples were collected immediately before and 15 min and 30 min after the formation of dominance relationships. Dominants showed an increase in plasma testosterone at 15 min post-interaction compared to their pre-interaction baseline, whereas subordinates and controls showed no change in plasma testosterone. Secondly, we investigated whether 14 days of dominant social status increased androgen or estrogen alpha-receptor immunoreactivity in brain regions that regulate the conditioned defeat response. Dominants showed more androgen, but not estrogen alpha, receptor immuno-positive cells in the dorsal medial amygdala (dMeA) and ventral lateral septum (vLS) compared to subordinates and controls. Finally, we showed that one day of dominant social status was insufficient to increase androgen receptor immunoreactivity compared to subordinates. These results suggest that elevated testosterone signaling at androgen receptors in the dMeA and vLS might contribute to the reduced conditioned defeat response exhibited by dominant hamsters.     

The effect of dominance on food hoarding: a game theoretical model

Many food hoarding animals live in small groups structured by rank. The presence of conspecifics in the hoarding area increases the risk of losing stored supplies. The possibility of stealing from others depends on a forager's rank in the group. Highly ranked individuals can steal from subordinates and also protect their own caches. Since storing incurs both costs and benefits, the optimal hoarding investment will differ between individuals of different rank. In a game theoretical model, we investigate how dominant and subordinate individuals should optimize their hoarding effort. Our model imagines animals that are large-scale hoarders in autumn and dependent on stored supplies for winter survival. Many examples can be found in the bird families Paridae and Corvidae, but the model can be used for any hoarding species that forage in groups. Predictions from the model are as follows: First, subordinates should store more than dominants, but in a predictable environment, this difference will decrease as the environment gets harsher. Under harsh conditions, dominants should store almost as much as subordinates and, later, spend almost as much time retrieving their own caches as subordinates. Second, if on the other hand, bad winter conditions were not expected when storing, dominants should spend more time pilfering caches from subordinates. Third, in populations that are highly dependent on stored supplies, dominants should store relatively more than in populations that are less dependent on stored supplies. Fourth, harsher environments will favor hoarding. And finally, if dominant individuals store, it implies that hoarders have a selfish recovery advantage over conspecific pilferers.     

Physical predictors,behavioural/emotional attributes and neurochemical determinants of dominant behaviour

Significant differences in physical and behavioural/emotional/cognitive predictors and attributes, as well as of neurochemical inducers of behaviour, between dominant and subordinate animals are discussed. It is still unknown whether these factors are the causes of differences between dominants and subordinates, or vice versa whether the differences between dominants and subordinates are the origin of differences in these factors. The possibility is discussed that no differences exist among juveniles in the concentrations of neurochemical agents (known in the literature as determinants of dominance) between the brains of future dominants and future subordinates. We describe a study design that makes the assessment of the ‘original’ neurochemical profile of the brain possible.     

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.     

Post-contest behaviour in black-capped chickadees (<Emphasis Type="Italic">Poecile atricapillus</Emphasis>): loser displays,not victory displays,follow asymmetrical countersinging exchanges

Victory displays are behaviours that occur after the conclusion of a signaling contest, performed solely by the contest winner. Victory displays may reinforce the dominance of the winner either to the loser or to other conspecifics within signaling range. Victory displays are poorly studied despite the significant consequences that post-conflict behaviour may have on the individuals involved. We examined the period immediately following 50 territorial countersinging contests between males in 10 neighbourhoods of black-capped chickadees (Poecile atricapillus) of known dominance rank. We characterized the post-contest singing behaviour of chickadees and evaluated whether post-contest behaviour is consistent with victory displays. Using a 16-microphone acoustic location system to simultaneously record entire neighbourhoods of breeding chickadees, we isolated 50 dyadic countersinging contests and measured the vocal behaviour of the contestants in the minutes following each interaction. Eighty-six percent of contests were followed by a period of solo singing by one of the contestants, while 14% were followed by silence. The post-contest singer was most often the contestant who held a subordinate dominance position in the previous winter’s dominance hierarchy; dominant males performed post-contest song bouts significantly less often. Asymmetry in overlapping between contestants did not predict which bird sang a post-contest bout. However, in a significant majority of cases, the post-contest singer was pitch-matched by his opponent during the contest more than he pitch-matched his opponent. Our results indicate that male chickadees do not perform acoustic victory displays after countersinging contests. In contrast, the post-contest behaviour of territorial chickadees is more consistent with a “loser display”.     

Differential response to the odor of familiar intruder mice in male mice (Mus musculus)

Responses to the odor of familiar intruder male mice according to their dominance were investigated. Responses were classified into 2 aspects: the investigation of the odor and the decision-making regarding avoiding it or not. The results varied according to the dominance of the respondents and the odor donor, and also according to the context of previous encounter situations. The dominants that had attacked an intruder dominant mouse responded randomly to its odor, whereas the dominants that had fought with it tended to avoid the odor. The subordinates that had observed an intruder dominant mouse being attacked by its dominant cagemate preferred the passage with the intruder's odor. The odor of a subordinate mouse was neither avoided nor preferred by either the dominants or subordinates. It was suggested that mice distinguished the dominance of the odor donor regardless of the context of the previous encounter situation, but they responded differently according to it and also according to their own dominance status.     

When should signals of submission be given?-A game theory model

During contests, losing animals often show signals communicating submission. From an evolutionary viewpoint, however, it is not so obvious why the losing individual gives such a signal instead of running away and why the winning individual accepts the signal instead of inflicting more severe damage. We investigated factors influencing the evolution of signals of submission using a numerical ESS model. The present analysis reveals that there is much space for the evolution of signals of submission, even when the winner of an escalated contest gets some extra benefit. In most cases, signals of submission are given by animals which are slightly or moderately weaker than the opponent. Signals of submission are expected to occur frequently (1) when the value of contested resource does not differ greatly from the cost of injury, (2) when the extra benefit of winning an escalated contest is small, (3) when the opportunity for safe retreat by the losing animal is small, and (4) when the estimation of the difference in the resource holding potential (RHP) between the combatants is accurate but not perfect.     

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.     

Intraspecific variation in competitive ability and food intake in salmonids: consequences for energy budgets and growth rates

The interactions between dominance status, feeding rate and growth in rainbow trout, Salmo gairdneri Richardson, were analyzed using published data on experimental populations. There was a positive correlation between metabolic expenditure and food intake in both dominant and subordinate fish, but dominants obtained a greater intake for a given expenditure than did subordinates. Subordinates that adopted a high–return/high–cost foraging strategy actually expended more energy than they acquired, whereas those that minimized energy expenditure obtained a net energy gain. This led to the surprising finding that the growth rate of subordinates was negatively correlated with food intake.     

Rank and reproduction in cooperatively breeding African wild dogs: behavioral and endocrine correlates

African wild dogs (Lycaon pictus) live in cooperative packswith a clear-cut dominance hierarchy in each sex. Reproductionis largely monopolized by the dominant male and female. Alphafemales produced 76% of all litters in the Selous Game Reserveand 81% in Kruger National Park. Only 6-17% of subordinate femalesgave birth each year, compared to 82% of dominant females. Innonmating periods, subordinate females had higher estrogen levelsand higher estrogen/progestin ratios than alpha females, apparentlypreventing ovulation. During mating periods, subordinate femaleshad lower estrogen levels than dominants, mated less often,and were less aggressive. Subordinate males mated at low rates,wore less aggressive than dominants, and had lower testosteronelevels. Beta males were similar to alpha males behavioraDy andhormonaUy, suggesting that alpha males may share paternity withbeta males. If paternity is more evenly shared than maternity,then subordinate males have a larger incentive than subordinatefemales to remain in the pack. Following this expectation, dispersalin Selous was female biased (49% versus 24% dispersing annually).Perhaps as a result of mortality associated with dispersal,the adult sex ratio was male biased, although the pup sex ratiowas unbiased. In Kruger, neither dispersal nor the adult sex-ratiowas biased. Reproductive suppression is widely thought to becaused by social stress in subordinates, but basal cortkosteronelevels were higher in dominants than in subordinates