Territorial aggression does not feed back on testosterone in a multiple-brooded songbird species with breeding and non-breeding season territoriality,the European stonechat

Testosterone mediates reproductive behaviours in male vertebrates. For example, breeding season territoriality depends on testosterone in many species of birds and in some, territorial interactions feed back on testosterone concentrations. However, the degree to which territorial behaviour and testosterone are associated differs even between species with seemingly similar life histories, especially between species that also defend territories outside the breeding season. Here, we investigate the link between territorial behaviour and testosterone in European stonechats. Previous studies found that territorial aggression in stonechats depends on testosterone in a breeding, but not in a non-breeding context. We investigated whether stonechats show a rise in testosterone during simulated territorial intrusions (STI) during the breeding season. Post-capture testosterone concentrations of males caught after an STI were not higher than those of males caught in a control situation regardless of breeding stage. However, most of the males would have been able to mount a testosterone response because the same individuals that did not increase testosterone during the STI showed a substantial increase in testosterone after injections of gonadotropin-releasing hormone (GnRH). GnRH-induced and post-capture testosterone concentrations were positively correlated and both decreased with successive breeding stages. Further, territory owners with a short latency to attack the decoy expressed higher post-capture testosterone concentrations than males with a longer latency to attack the decoy. Thus, there is no evidence for behavioural feedback on testosterone concentrations during male-male interactions in stonechats. In combination with previous studies our data suggest that testosterone functions as an on/off switch of high intensity territorial aggression during the breeding season in stonechats. The among-species variation in the androgen control of territorial behaviour may be only partly a result of environmental differences. Instead, potential differences in how territoriality evolved in different species may have influenced whether and how a reproductive hormone such as testosterone was co-opted into the mechanistic control of territorial behaviour.     

Social Experience During Development and Female Offspring Aggression in Peromyscus Mice

Cross‐fostering between the highly aggressive, biparental California mouse (Peromyscus californicus) and the less aggressive, less parental white‐footed mouse (P. leucopus) influences female offspring attack latency in California mice, but not in white‐footed mice. Adult female California mice raised by white‐footed mice expressed longer attack latencies in a neutral‐arena test but not in a resident‐intruder test. One social cue that may be used by offspring to develop environmentally appropriate levels of aggression is the type of parental care during development. In California mice, a composite score of maternal behavior was positively associated with neutral‐arena aggression as indicated by decreased attack latency. In both species, paternal nest‐building was positively associated with neutral‐arena aggression and higher maternal retrieval behavior predicted higher offspring resident‐intruder aggression as indicated by decreased attack latency. Together, these results indicate that parental behavior has the potential to shape the development of attack latency in female offspring.     

Differences in basal and stress-induced HPA regulation of wild house mice selected for high and low aggression

Male wild house mice, selected for short (SAL) and long (LAL) attack latency, show distinctly different behavioral strategies in coping with environmental challenges. In this study, we tested the hypothesis that this difference in coping style is associated with a differential stress responsiveness of the hypothalamic-pituitary-adrenal (HPA) system. SAL rather than LAL mice showed a clear fluctuation in circulating corticosterone concentrations around the circadian peak with significantly higher levels in the late light phase. LAL mice showed lower basal ACTH levels and higher thymic and spleen weights compared to SAL. Under basal conditions, glucocorticoid receptor (GR) and mineralocorticoid receptor (MR) mRNA in the hippocampus and corticotropin-releasing hormone (CRH) mRNA in the paraventricular nucleus of the hypothalamus were not different between the two lines. Forced swimming for 5 min induced high immobility behavior in LAL mice which was associated with an enhanced and prolonged corticosterone response as compared to SAL, while absolute ACTH levels did not differ. In addition, LAL mice showed an increase in hippocampal MR mRNA (but not GR) and hypothalamic CRH mRNA at 24 h after forced swimming. In conclusion, a genetic trait in coping style of wild house mice is associated with an idiosyncratic pattern of HPA activity, and greater responsiveness of physiological and molecular stress markers in LAL mice. In view of the profound differences in behavioral traits and stress system reactivity, these mouse lines genetically selected for attack latency present an interesting model for studying the mechanism underlying individual variation in susceptibility to stress-related psychopathology.     

Acute changes in plasma testosterone levels and their relation to measures of sexual behaviour in the male house mouse (Mus musculus)

There is an increasing amount of evidence, from several mammalian species, that plasma testosterone levels increase acutely when the male is exposed to sexual stimuli. Work reported here investigates whether such responses occur in male mice, and whether their incidence is related to measures of sexual behaviour. Close proximity of oestrous female mice was found to result in increased plasma testosterone levels in some strains within 15 min. Immediately after a sexual behaviour test, plasma testosterone levels were higher in males showing sexual responses. There were correlations between testosterone levels and mount latency, but none with ejaculation latency. Testosterone levels were greatest at the initiation of mounting responses, and declined during copulation, although not significantly.     

Individual differences in the attack behavior of male mice: a function of attack stimulus and hormonal state

Male CFW mice were tested for fighting behavior directed against olfactory bulbectomized male mice and against lactating female mice. Some males were tested with each stimulus type before and after castration. Some males were tested first following castration and then after testosterone treatment. All gonadally intact males attacked bulbectomized males and 25% attacked lactating females. After castration 81% attacked males on at least one occasion and 62% began to attack lactating females, although individual differences in the pattern of post-castration behavior were large. Individual differences in attack behavior were also large in males whose first tests followed castration. Of these, 60% attacked both stimuli. Following testosterone treatment, attack against males increased while attack against females was inhibited. Hormonal stimulation reduced individual differences in behavior and increased males' discrimination between the two types of stimuli.     

Predatory aggression, but not maternal or intermale aggression, is associated with high voluntary wheel-running behavior in mice

Predatory (towards crickets), intermale, and maternal aggression were examined in four replicate lines of mice that had been selectively bred for high wheel-running (S) and in four random-bred control lines (C). In generation 18, individual differences in both predatory and intermale aggression were significantly consistent across four trial days, but predatory and intermale aggression were uncorrelated both at the individual level and among the eight line means. Latencies to attack crickets were significantly lower in S lines as a group. Intermale aggression, however, did not differ between S and C lines. S lines were significantly smaller in body mass, but did not differ in either testes mass or plasma testosterone. In generations 28 and 30, respectively, S and C lines did not differ in either maternal or intermale aggression. However, significant differences among the individual lines were found for maternal aggression, and one S line exhibited an extremely high mean time of aggression (>120 sec for a 5-min test). Maternal and intermale aggression were not correlated among the eight line means or at the level of individual variation. Overall, our results suggest: (1) predatory aggression and voluntary wheel-running are positively related at the genetic level; (2) predatory and intermale aggression are unrelated at a genetic level; and (3) maternal and intermale aggression are not tightly related at the genetic level. Possible relationships between predatory aggression, dopamine, and wheel-running behavior are discussed.     

Age-associated changes in plasma testosterone levels in male mice and their relation to social dominance or subordinance

Plasma testosterone (T) levels were determined in male mice of the CD-1 (ICR) strain from 30 to 680 days of age. The mean plasma T was 5.2 ± 1.0 ng/ml for adult male mice (70–400 days old, 102 mice) and 1.8 ± 0.9 ng/ml for old males (450–680 days old, 11 mice). Through 140 determinations, the highest T value obtained was 52.3 ng/ml in an 80-day-old individual while the lowest was 0.08 ng/ml in a 680-day-old animal. It was found that plasma T was significantly decreased in the old age, although individual variation in T levels was considerable. On the other hand, social dominance or subordinance was examined among male mice in each cage and its relation to plasma T was investigated. Dominance or subordinance was verified by frequency of chases or attacks delivered or received, or number of fights won or lost. It was observed that dominant-subordinate relationships were present in 6 out of 10 cages and that the dominant individual had a significantly higher T level than the subordinate male; the mean T level was 10.5 ± 2.5 ng/ml for the dominant individuals vs 2.2 ± 0.8 ng/ml for the subordinate ones. Marked individual variation in plasma T titers in male mice seems to be related to the dominance/subordinance rank within a group.     

Serotonin Depletion-Induced Maladaptive Aggression Requires the Presence of Androgens

The sex hormone testosterone and the neurotransmitter serotonin exert opposite effects on several aspects of behavior including territorial aggression. It is however not settled if testosterone exerts its pro-aggressive effects by reducing serotonin transmission and/or if the anti-aggressive effect of serotonin requires the presence of the androgen. Using the resident intruder test, we now show that administration of the serotonin synthesis inhibitor para-chlorophenylalanine (300 mg/kg x 3 days) increases the total time of attack as well as the percentage amount of social behavior spent on attack but not that spent on threat – i.e. that it induces a pattern of unrestricted, maladaptive aggression – in gonadectomized C57Bl/6 male mice receiving testosterone replacement; in contrast, it failed to reinstate aggression in those not given testosterone. Whereas these results suggest the pro-aggressive effect of testosterone to be independent of serotonin, and not caused by an inhibition of serotonergic activity, the pCPA-induced induction of maladaptive aggression appears to require the presence of the hormone. In line with these findings, pCPA enhanced the total time of attack as well the relative time spent on attacks but not threats also in wild-type gonadally intact male C57Bl/6 mice, but failed to reinstate aggression in mice rendered hypo-aggressive by early knock-out of androgen receptors in the brain (AR^NesDel mice). We conclude that androgenic deficiency does not dampen aggression by unleashing an anti-aggressive serotonergic influence; instead serotonin seems to modulate aggressive behavior by exerting a parallel-coupled inhibitory role on androgen-driven aggression, which is irrelevant in the absence of the hormone, and the arresting of which leads to enhanced maladaptive aggression.     

Intensified aggressivity of dominant male mice after the intraventricular administration of 6-hydroxydopamine

Aggressiveness of dominant male mice of different genotypes was studied after administration of 6-OHDA into the brain lateral ventriculus. Administration of neurotoxine led to lowering of noradrenaline and dopamine level in the brain in the average by 40%. Besides, the aggressiveness of the animals rose and the latency of the first attack shortened, while their rank did not change. Administration of the solvent in analogous conditions influenced neither the rank of the animal nor its aggressiveness. Conclusion is made that catecholamines take part in regulation of aggressiveness of dominant male mice.     

Interplay of individual interactions and turbidity affects the functional response of three-spined sticklebacks Gasterosteus aculeatus

The effects of turbidity, size and the presence of conspecifics on the functional response, feeding latency and activity in the three-spined stickleback Gasterosteus aculeatus were examined. A significant interaction between standard length and presence of conspecifics demonstrated an increase in attack rates of larger individuals in the presence of conspecifics. Attack rate was also higher in turbid water. Feeding latency decreased with prey concentration and presence of conspecifics, but was not affected by turbidity. Activity level did not change with prey levels, but increased with turbidity. These results can help to better understand how individual flexibility in the functional response can affect prey mortality according to environmental perturbation and social interaction at the level of the predator.     

Crowding pregnant mice affects attack and threat behavior of male offspring

Attack and threat behavior of adult male offspring of female mice crowded during the final third of pregnancy was investigated. In 5-min test pairings with an anosmic "standard opponent" which had 50 microliter of male mouse urine applied to its fur, the prenatally stressed group of males showed significantly less attack behavior; attack latency was longer and number of attacks, bites, amount of time spent attacking, and composite aggression scores were all lower, compared with the control group. Similarly, less threat behavior was observed in offspring from crowded dams; there were lower frequencies of tail rattles, rough grooms, and upright threats. Additionally, proportionally fewer males in the prenatally stressed group attacked or displayed threats. A second experiment was designed to investigate the effects of exogenous androgen on the aggressiveness of males from crowded mice: testosterone propionate administration (500 micrograms/animal/day, for 5 days prior to testing) abolished differences both in the proportion of males from crowded mice that fought and also apparently abolished differences in intensities of attack and threat behavior between groups. However, trends toward reduced aggression in prenatally crowded males remained. More detailed analysis of these responses, based only on animals that displayed aggression, revealed significantly reduced intensity of aggression in offspring from females crowded during pregnancy, indicating that testosterone propionate therapy did not completely restore this behavior. In order to reduce postnatal effects due to possible differences in mothering, all offspring were fostered to untreated mothers at birth. The results are discussed in terms of in utero exposure of male fetuses of crowded dams to stress-liberated adrenal steroids of maternal origin, and the possible consequences for the endocrine integrity of these offspring.     

The Use of Standard Aggression Testing Methods to Predict Combat Behaviour and Contest Outcome in Rivulus marmoratus Dyads (Teleostei: Cyprinodontidae)

Aggression plays an important role in animal contests, but the extent to which aggression correlates with dominance has been a topic of much debate. The relationship between aggression and dominance ability in the hermaphroditic fish, Rivulus marmoratus , was investigated using three standard protocols, the mirror test (Mi), model test (Mo), and standard opponent test (So). In each, display latency, attack latency, and biting frequency were quantified for a test individual towards its opponent. The general rank-order for eliciting strength of the three different stimuli was Mi > So > Mo. The relationships between the individual indices from the standard tests and three dyadic contest variables, initiator of display, initiator of attack, and winner, were analysed in contests between previously tested pairs to ascertain how well the standard protocols predicted dyadic contest behaviour/outcome. Display and attack latencies in the standard tests did not predict the level of analogous combat behaviour. Biting frequency differences between individuals in a pair in the So and Mo tests as well as display latency differences in the Mi test contributed to predictions of contest outcome. The individual that scored higher, relative to its opponent, won a significantly greater proportion of the bouts. These findings demonstrate the importance of relative differences in aggression in determining dominance. However, the predictive value of standard test behaviour is test-specific and, based on the available literature, depends on both the species used and the context in which they are employed.     

Androgen influence on male mouse ultrasounds during courtship

Ultrasonic vocalizations are emitted by adult male mice (Mus musculus) shortly after an adult female or her odor are encountered, possibly as an early part of courtship behavior. Consistent with this hypothesis, it was found that castration of adult male mice increased the latency to first ultrasonic vocalization in response to an adult female. In addition, castrated males, subsequently injected once with 200 μg of testosterone propionate, reduced their latency, whereas oil-injected castrates did not.     

Endogenous testosterone levels do not affect filarial worm burdens in mice.

It has been observed for several years that males of many mammalian species exhibit higher prevalence and intensity of parasitic infections than females. It has been theorized that this sexual dichotomy may be due to the immunosuppressive properties of testosterone. The administration of exogenous testosterone to laboratory animals often results in higher yields of several parasitic nematodes, lending credence to this hypothesis. We sought to determine if worm burdens in individual mice were related to serum testosterone concentrations. We found that in a cage of five male mice, one or two individuals exhibited high circulating testosterone levels, while the rest had low to undetectable levels. Serum testosterone did not correlate with worm recoveries. The implications of these findings are twofold. First, our data suggest that high endogenous testosterone does not necessarily decrease the ability of mice to resist nematode infection and second, that results obtained from administration of exogenous testosterone should be interpreted with caution.     

Anticipated,Intense Risk Enhances Behavioral Conformity in a Mouse Model

A mouse conformity model was established by contrasting the differences in the time spent navigating a custom‐made maze when mice were alone versus when they were in a companion group. Experimental mice received one‐trial inhibitory avoidance training by experiencing a foot shock after they arrived at a designated goal box in the maze. In contrast, four mice of the respective companion group received food pellet trainings for rapidly approaching the same goal box. In the retest, mice receiving 0.3‐mA shock in the training trial and navigating alone displayed longer arrival latency to the goal box compared with the mice receiving same level of shock in the training trial and navigating along with the companion group. A conformity index was calculated for the arrival latency of the experimental mice receiving the retest alone and the latency of the experimental mice receiving the retest along with the companion group. In the retest, the conformity index progressively increased with the training foot shock level. No difference was noticed in the arrival latency to the goal box or the conformity index between group‐navigating experimental mice accompanied by familiar and unfamiliar groups. These results, taken together, indicate that behavioral conformity can be enhanced by an expected, intense risk. Moreover, familiarity with the companion group does not affect the behavioral conformity of an individual. We conclude that species members‐shared risk‐detecting and risk‐coping systems could be critical in motivating behavioral conformity for each member of this species.     

Does testosterone affect foraging behavior in male frogs?

During the breeding season, males often produce costly and extravagant displays or physical ornaments to attract females. Numerous studies have established that testosterone could directly influence the expression of certain sexual signals. However, few of these studies have focused on the indirect role that testosterone could play in modulating prey detection and visual performance to improve the foraging ability of males and hence their acquisition of nutritional resource. In the present study, we experimentally modified the testosterone levels of European tree frog males (Hyla arborea), staying in the natural range previously measured in the field, and we investigated the effect of testosterone on the foraging ability of individuals. Foraging capacities were measured on males placed in an arena with a virtual cricket moving on a computer screen. Our results demonstrated a significant effect of testosterone on the hunting behavior of H. arborea. We observed that testosterone reduced the orientation latency to virtual prey for supplemented males compared to controls. In addition, testosterone significantly increased the attack promptness of male frogs. Finally, our experiment did not demonstrate any impact of testosterone on male attack success.     

Aggressive and non-aggressive personalities differ in oxidative status in selected lines of mice (Mus musculus)

Mice selected for aggression and coping (long attack latency (LAL), reactive coping strategy; short attack latency (SAL), pro-active coping strategy) are a useful model for studying the physiological background of animal personalities. These mice also show a differential stress responsiveness, especially in terms of hypothalamic-pituitary-adrenal axis reactivity, to various challenges. Since the stress response can increase the production of reactive oxygen species, we predicted that the basic oxidative status of the lines could differ. We found that LAL showed higher serum antioxidant capacity (OXY) than SAL, while no differences emerged for reactive oxygen metabolites (ROMs) or the balance between ROMs and OXY, reflecting oxidative stress. Moreover, the lines showed inverse relationships between ROMs or OXY and body mass corrected for age. The results indicate that variation in oxidative status is heritable and linked to personality. This suggests that different animal personalities may be accompanied by differences in oxidative status, which may predict differences in longevity.     

Evidence for episodic secretion of testosterone in laboratory mice.

The concentration of testosterone (T) in the peripheral plasma of laboratory mice is extremely variable. This variability is already evident at 20--25 days of age and is not eliminated by brief or chronic exposure to male or female mice, or by isolation. The variation in T levels in plasma samples collected from the same animals on different occasions is comparable to the variation between individuals bled on a single occasion. The concentration of T in the testis is as variable as that in the peripheral plasma. It is suggested that in the laboratory mouse T is produced and released in an episodic fashion, that elevations in T levels in peripheral plasma of mice are greater than those observed in other species, and that testicular secretory episodes are interspersed with periods of minimal steroidogenic activity.     

Comparative characteristics of the parameters of an aggressive reaction in 2 mouse genotypes

The parameters have been studied of the aggressive reaction of male mice of CBA/Lac and C57BL/6J lines differing by olfactory sensitivity to zoosocial pheromone stimuli. It has been shown that CBA males, characterized by a high olfactory sensitivity, have lower latency of the first attack than C57BL males with low olfactory sensitivity. A prolonged distant exposition to an unknown litter and male appearance lowers the latency of the first attack in mice of the studied lines proportionally to their meanings demonstrated after short time exposition. The number of attacks and total time of attacking is considerably higher in C57BL mice during the whole test period (15 min) than in CBA mice in which aggressivity is already sharply lowered after 5 min of agonistic interactions. The factors are discussed, influencing the parameters of mice aggressive reaction.