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1.  Altered Immunity in Crowded Locust Reduced Fungal (Metarhizium anisopliae) Pathogenesis 
PLoS Pathogens  2013;9(1):e1003102.
The stress of living conditions, similar to infections, alters animal immunity. High population density is empirically considered to induce prophylactic immunity to reduce the infection risk, which was challenged by a model of low connectivity between infectious and susceptible individuals in crowded animals. The migratory locust, which exhibits polyphenism through gregarious and solitary phases in response to population density and displays different resistance to fungal biopesticide (Metarhizium anisopliae), was used to observe the prophylactic immunity of crowded animals. We applied an RNA-sequencing assay to investigate differential expression in fat body samples of gregarious and solitary locusts before and after infection. Solitary locusts devoted at least twice the number of genes for combating M. anisopliae infection than gregarious locusts. The transcription of immune molecules such as pattern recognition proteins, protease inhibitors, and anti-oxidation proteins, was increased in prophylactic immunity of gregarious locusts. The differentially expressed transcripts reducing gregarious locust susceptibility to M. anisopliae were confirmed at the transcriptional and translational level. Further investigation revealed that locust GNBP3 was susceptible to proteolysis while GNBP1, induced by M. anisopliae infection, resisted proteolysis. Silencing of gnbp3 by RNAi significantly shortened the life span of gregarious locusts but not solitary locusts. By contrast, gnbp1 silencing did not affect the life span of both gregarious and solitary locusts after M. anisopliae infection. Thus, the GNBP3-dependent immune responses were involved in the phenotypic resistance of gregarious locusts to fungal infection, but were redundant in solitary locusts. Our results indicated that gregarious locusts prophylactically activated upstream modulators of immune cascades rather than downstream effectors, preferring to quarantine rather than eliminate pathogens to conserve energy meanwhile increasing the “distance” of infectious and target individuals. Our study has obvious implications for bio-pesticides management of crowded pests, and for understanding disease epidemics and adaptiveness of pathogens.
Author Summary
The wide application of fungal biopesticides for insect management has led to concerns over the development of biopesticide resistance. The migratory locust, a globally notorious agricultural pest, has density-dependent phase changes between solitary and gregarious states. The gregarious locusts displayed longer life spans than solitary locusts after biopesticide Metarhizium anisopliae infection. We analyzed prophylactic immunity of the locusts in phase change adaptation by transcriptome analysis. Gregarious locusts optimized immunity by investing more in molecules of upstream immune cascades including pattern recognition proteins, anti-oxidation proteins, protease inhibitors and serine protease. High levels of pattern recognition proteins guided deposition of immune products onto pathogens reducing growth, proliferation and transmission. This prophylactic immunity of gregarious locusts emphasized on quarenteening M. anisopliae pathogens in early infection, which decreased individuals' infection risk in a population and avoids disease epidemics. Pest outbreaks mostly occur in high population densities, thereby, diminishing entomopathogen biopesticide efficiency. Our results provide an insight to an organism's “enhanced” immunity induced by population densities and inspires new paradigms to understand biopesticide tolerance and disease epidemics in the future.
doi:10.1371/journal.ppat.1003102
PMCID: PMC3542111  PMID: 23326229
2.  Social bonds and rank acquisition in raven nonbreeder aggregations 
Animal Behaviour  2012;84(6):1507-1515.
Complex social life has been characterized as cognitively challenging and recently, social relationships such as long-term social bonds and alliances have been identified as key elements for brain evolution. Whereas good evidence is available to support the link between social relations and cognition in mammals, it remains unsatisfying for birds. Here we investigated the role of avian social bonds in a nonbreeder aggregation of ravens, Corvus corax, in the Austrian Alps. We individually marked 138 wild ravens, representing approximately half of a population that uses the area of a local zoo for foraging. For 2 years, we observed the dynamics of group composition and the birds' agonistic and affiliative interactions. We identified two levels of organization: the formation of an unrelated local group and the individuals' engagement in social bonds of different length and reciprocity pattern. Whereas belonging to the local group had no significant effect on conflicts won during foraging, the individual bonding type did. Birds that engaged in affiliative relationships were more successful when competing for food than those without such bonds. Bonded birds did suffer from aggression by other bonded birds and, probably as a consequence, most of the ravens' social relations were not stable over time. These results support the idea that social bonding and selective cooperation and competition are prominent features in nonbreeding ravens. Proximately, bonding may qualify as a social manoeuvre that facilitates access to resources; ultimately it might function to assess the quality of a partner in these long-term monogamous birds.
Highlights
► Big brains correlate with the formation of strong nonsexual bonds in mammals. ► We examine formation and use of bonds in birds, namely wild nonbreeder ravens. ► Individuals engage in different social bonds and benefit from them in conflicts. ► Active investment in nonsexual strong bonds for rank acquisition in a big-brained bird.
doi:10.1016/j.anbehav.2012.09.024
PMCID: PMC3518779  PMID: 23264693
Corvus corax; dominance; nonbreeder aggregation; raven; social bond; social structure
3.  Rich Pickings Near Large Communal Roosts Favor ‘Gang’ Foraging by Juvenile Common Ravens, Corvus corax 
PLoS ONE  2009;4(2):e4530.
Ravens (Corvus corax) feed primarily on rich but ephemeral carcasses of large animals, which are usually defended by territorial pairs of adults. Non-breeding juveniles forage socially and aggregate in communal winter roosts, and these appear to function as ‘information centers’ regarding the location of the rare food bonanzas: individuals search independently of one another and pool their effort by recruiting each other at roosts. However, at a large raven roost in Newborough on Anglesey, North Wales, some juveniles have been observed recently to forage in ‘gangs’ and to roost separately from other birds. Here we adapt a general model of juvenile common raven foraging behavior where, in addition to the typical co-operative foraging strategy, such gang foraging behavior could be evolutionarily stable near winter raven roosts. We refocus the model on the conditions under which this newly documented, yet theoretically anticipated, gang-based foraging has been observed. In the process, we show formally how the trade off between search efficiency and social opportunity can account for the existence of the alternative social foraging tactics that have been observed in this species. This work serves to highlight a number of fruitful avenues for future research, both from a theoretical and empirical perspective.
doi:10.1371/journal.pone.0004530
PMCID: PMC2646834  PMID: 19240813
4.  Predators as stressors? Physiological and reproductive consequences of predation risk in tropical stonechats (Saxicola torquata axillaris). 
Tropical birds usually lay smaller clutches and are less likely to initiate a second brood than their temperate-zone relatives. This reduction in annual fecundity is generally explained as an adaptation either to higher rates of nest predation or to a more limited food supply concurrent with higher adult survival in the tropics. However, the physiological parameters associated with lower annual fecundity in tropical birds have not been well investigated. We compared the annual fecundity, behaviour and a number of physiological parameters of stonechat parents feeding fledged juveniles in territories with and without fiscal shrikes, a predator on adult and fledged birds. Stonechat pairs in territories with shrikes were less likely to initiate a second brood and delayed successive broods compared to pairs in territories without shrikes. After fledging of their young, males showed a greater propensity than females to initiate distraction calls after a human intrusion into their territory and, therefore, invested more in the defence of their young. In territories with shrikes stonechat males had higher initial plasma corticosterone levels and lower body conditions than males in territories without shrikes, suggesting that they were chronically stressed. In contrast, the females from both types of territory had low initial plasma corticosterone levels. We conclude that shrike presence might account for the delay in initiation of a second brood and the reduction in the tendency to initiate a second brood. Whether these effects are mediated by the elevated levels of corticosterone remains to be demonstrated.
doi:10.1098/rspb.2001.1691
PMCID: PMC1088780  PMID: 11487404
5.  Reciprocity of agonistic support in ravens 
Animal Behaviour  2012;83(1):171-177.
Cooperative behaviour through reciprocation or interchange of valuable services in primates has received considerable attention, especially regarding the timeframe of reciprocation and its ensuing cognitive implications. Much less, however, is known about reciprocity in other animals, particularly birds. We investigated patterns of agonistic support (defined as a third party intervening in an ongoing conflict to attack one of the conflict participants, thus supporting the other) in a group of 13 captive ravens, Corvus corax. We found support for long-term, but not short-term, reciprocation of agonistic support. Ravens were more likely to support individuals who preened them, kin and dominant group members. These results suggest that ravens do not reciprocate on a calculated tit-for-tat basis, but aid individuals from whom reciprocated support would be most useful and those with whom they share a good relationship. Additionally, dyadic levels of agonistic support and consolation (postconflict affiliation from a bystander to the victim) correlated strongly with each other, but we found no evidence to suggest that receiving agonistic support influences the victim’s likelihood of receiving support (consolation) after the conflict ends. Our findings are consistent with an emotionally mediated form of reciprocity in ravens and provide additional support for convergent cognitive evolution in birds and mammals.
Highlights
► We examine patterns of agonistic support in captive ravens. ► We find support for long-term but not short-term reciprocation of support. ► Ravens supported individuals who preened them, kin and dominant individuals. ► Strong correlation between dyadic levels of agonistic support and consolation. ► Ravens may use emotions to mediate reciprocation of support.
doi:10.1016/j.anbehav.2011.10.023
PMCID: PMC3255075  PMID: 22298910
agonistic support; coalition; corvid; Corvus corax; interchange; raven; reciprocity
6.  Corticosterone, testosterone and life-history strategies of birds 
Steroid hormones have similar functions across vertebrates, but circulating concentrations can vary dramatically among species. We examined the hypothesis that variation in titres of corticosterone (Cort) and testosterone (T) is related to life-history traits of avian species. We predicted that Cort would reach higher levels under stress in species with higher annual adult survival rates since Cort is thought to promote physiological and behavioural responses that reduce risk to the individual. Conversely, we predicted that peak T during the breeding season would be higher in short-lived species with high mating effort as this hormone is known to promote male fecundity traits. We quantified circulating hormone concentrations and key life-history traits (annual adult survival rate, breeding season length, body mass) in males of free-living bird species during the breeding season at a temperate site (northern USA) and a tropical site (central Panama). We analysed our original data by themselves, and also combined with published data on passerine birds to enhance sample size. In both approaches, variation in baseline Cort (Cort0) among species was inversely related to breeding season length and body mass. Stress-induced corticosterone (MaxCort) also varied inversely with body mass and, as predicted, also varied positively with annual adult survival rates. Furthermore, species from drier and colder environments exhibited lower MaxCort than mesic and tropical species; T was lowest in species from tropical environments. These findings suggest that Cort0, MaxCort and T modulate key vertebrate life-history responses to the environment, with Cort0 supporting energetically demanding processes, MaxCort promoting survival and T being related to mating success.
doi:10.1098/rspb.2010.0673
PMCID: PMC2982062  PMID: 20554550
life-history trade-offs; steroid hormone; latitude; bird; reproduction; survival
7.  Loss of density-dependence and incomplete control by dominant breeders in a territorial species with density outbreaks 
BMC Ecology  2011;11:16.
Background
A territory as a prerequisite for breeding limits the maximum number of breeders in a given area, and thus lowers the proportion of breeders if population size increases. However, some territorially breeding animals can have dramatic density fluctuations and little is known about the change from density-dependent processes to density-independence of breeding during a population increase or an outbreak. We suggest that territoriality, breeding suppression and its break-down can be understood with an incomplete-control model, developed for social breeders and social suppression.
Results
We studied density dependence in an arvicoline species, the bank vole, known as a territorial breeder with cyclic and non-cyclic density fluctuations and periodically high densities in different parts of its range. Our long-term data base from 38 experimental populations in large enclosures in boreal grassland confirms that breeding rates are density-regulated at moderate densities, probably by social suppression of subordinate potential breeders. We conducted an experiment, were we doubled and tripled this moderate density under otherwise the same conditions and measured space use, mortality, reproduction and faecal stress hormone levels (FGM) of adult females. We found that mortality did not differ among the densities, but the regulation of the breeding rate broke down: at double and triple densities all females were breeding, while at the low density the breeding rate was regulated as observed before. Spatial overlap among females increased with density, while a minimum territory size was maintained. Mean stress hormone levels were higher in double and triple densities than at moderate density.
Conclusions
At low and moderate densities, breeding suppression by the dominant breeders, But above a density-threshold (similar to a competition point), the dominance of breeders could not be sustained (incomplete control). In our experiment, this point was reached after territories could not shrink any further, while the number of intruders continued to increase with increasing density. Probably suppression becomes too costly for the dominants, and increasing number of other breeders reduces the effectiveness of threats. In wild populations, crossing this threshold would allow for a rapid density increase or population outbreaks, enabling territorial species to escape density-dependency.
doi:10.1186/1472-6785-11-16
PMCID: PMC3141362  PMID: 21726437
8.  Aggressive Transition between Alternative Male Social Tactics in a Long-Lived Australian Dragon (Physignathus lesueurii) Living at High Density 
PLoS ONE  2012;7(8):e41819.
Theory predicts the evolution of alternative male social tactics when intense competition coupled with the superior competitive ability of some individuals limits access to reproductive opportunities by others. How selection has shaped alternative social tactics may be especially interesting in long-lived species where size among sexually mature males varies markedly. We conducted experimental studies on long-lived eastern Australian water dragons living where competition was intense to test the hypotheses that mature males adopt alternative social tactics that are plastic, and that large size and body condition determine resource-holding potential. Approximately one-half of mature males (N = 14) defended territories using high rates of patrol and advertisement display, whereas 16 smaller mature males having lower body condition indices utilized non-territorial social tactics. Although territorial males were larger in absolute size and head dimensions, their heads were not allometrically larger. Territorial males advertised very frequently using displays involving stereotypical movements of the head and dewlap. More aggressive displays were given infrequently during baseline social conditions, but increased during periods of social instability. Female home ranges overlapped those of several territorial and non-territorial males, but females interacted more frequently with territorial males. The extreme plasticity of social tactics in this species that are dependent on body size was confirmed by two instances when relatively large non-territorial males spontaneously evicted territory owners, and by marked shifts in tactics by non-territorial males in response to temporary experimental removals of territory owners, followed (usually) by their expulsion when original owners were reinstated. The high level of social plasticity in this population where same-sex competitors are densely concentrated in preferred habitat suggests that chronic high energetic costs of defense may select for males to cycle between territorial and non-territorial social tactics depending upon their changing energetic status and their current capacity for competition with rivals.
doi:10.1371/journal.pone.0041819
PMCID: PMC3414507  PMID: 22905109
9.  Long-Term Memory for Affiliates in Ravens 
Current Biology  2012;22(9):801-806.
Summary
Complex social life requires individuals to recognize and remember group members [1] and, within those, to distinguish affiliates from nonaffiliates. Whereas long-term individual recognition has been demonstrated in some nonhuman animals [2–5], memory for the relationship valence to former group members has received little attention. Here we show that adult, pair-housed ravens not only respond differently to the playback of calls from previous group members and unfamiliar conspecifics but also discriminate between familiar birds according to the relationship valence they had to those subjects up to three years ago as subadult nonbreeders. The birds' distinction between familiar and unfamiliar individuals is reflected mainly in the number of calls, whereas their differentiation according to relationship valence is reflected in call modulation only. As compared to their response to affiliates, ravens responded to nonaffiliates by increasing chaotic parts of the vocalization and lowering formant spacing, potentially exaggerating the perceived impression of body size. Our findings indicate that ravens remember relationship qualities to former group members even after long periods of separation, confirming that their sophisticated social knowledge as nonbreeders is maintained into the territorial breeding stage.
Highlights
► Ravens remember former group members for extended time periods ► Ravens memorize relationship valence (affiliation) in addition to group membership ► Familiarity is coded in the number of calls given in response to playback ► Relationship valence is coded in the modulation of response calls
doi:10.1016/j.cub.2012.03.023
PMCID: PMC3348500  PMID: 22521788
10.  High Risk of Lead Contamination for Scavengers in an Area with High Moose Hunting Success 
PLoS ONE  2014;9(11):e111546.
Top predators and scavengers are vulnerable to pollutants, particularly those accumulated along the food chain. Lead accumulation can induce severe disorders and alter survival both in mammals (including humans) and in birds. A potential source of lead poisoning in wild animals, and especially in scavengers, results from the consumption of ammunition residues in the tissues of big game killed by hunters. For two consecutive years we quantified the level lead exposure in individuals of a sentinel scavenger species, the common raven (Corvus corax), captured during the moose (Alces alces) hunting season in eastern Quebec, Canada. The source of the lead contamination was also determined using stable isotope analyses. Finally, we identified the different scavenger species that could potentially be exposed to lead by installing automatic cameras targeting moose gut piles. Blood lead concentration in ravens increased over time, indicating lead accumulation over the moose-hunting season. Using a contamination threshold of 100 µg.L−1, more than 50% of individuals were lead-contaminated during the moose hunting period. Lead concentration was twice as high in one year compared to the other, matching the number of rifle-shot moose in the area. Non-contaminated birds exhibited no ammunition isotope signatures. The isotope signature of the lead detected in contaminated ravens tended towards the signature from lead ammunition. We also found that black bears (Ursus americanus), golden eagles and bald eagles (Aquila chrysaetos and Haliaeetus leucocephalus, two species of conservation concern) scavenged heavily on moose viscera left by hunters. Our unequivocal results agree with other studies and further motivate the use of non-toxic ammunition for big game hunting.
doi:10.1371/journal.pone.0111546
PMCID: PMC4229082  PMID: 25389754
11.  Progesterone modulates aggression in sex-role reversed female African black coucals 
Testosterone is assumed to be the key hormone related to resource-defence aggression. While this role has been confirmed mostly in the context of reproduction in male vertebrates, the effect of testosterone on the expression of resource-defence aggression in female vertebrates is not so well established. Furthermore, laboratory work suggests that progesterone inhibits aggressive behaviour in females. In this study, we investigated the hormonal changes underlying territorial aggression in free-living female African black coucals, Centropus grillii (Aves; Cuculidae). Females of this sex-role reversed polyandrous bird species should be particularly prone to be affected by testosterone because they aggressively defend territories similar to males of other species. We show, however, that territorial aggression in female black coucals is modulated by progesterone. After aggressive territorial challenges female black coucals expressed lower levels of progesterone than unchallenged territorial females and females without territories, suggesting that progesterone may suppress territorial aggression and is downregulated during aggressive encounters. Indeed, females treated with physiological concentrations of progesterone were less aggressive than females with placebo implants. This is one of the first demonstrations of a corresponding hormone–behaviour interaction under challenged and experimental conditions in free-living females. We anticipate that our observation in a sex-role reversed species may provide a more general mechanism, by which progesterone—in interaction with testosterone—may regulate resource-defence aggression in female vertebrates.
doi:10.1098/rspb.2007.1707
PMCID: PMC2600909  PMID: 18252672
territorial aggression; females; testosterone; progesterone; sex-role reversal; classical polyandry
12.  Modulation of the hypothalamic-pituitary-adrenal axis of an Arctic-breeding polygynandrous songbird, the Smith's longspur, Calcarius pictus. 
To successfully reproduce in the Arctic, birds must modulate their neuroendocrine and behavioural systems. These adjustments include an attenuation of the stress responsiveness of the hypothalamic-pituitary-adrenal (HPA) axis to external stimuli and a behavioural insensitivity to high corticosterone (B) levels. The HPA axis was examined in free-living territorial polygynandrous Smith's longspurs (Calcarius pictus) that migrate to breed on the Arctic tundra. Basal and stress-induced B levels were measured through the breeding season and were found to be significantly lower in females compared with males. This was not a consequence of adrenal insensitivity, because intrajugular injections of adrenocorticotrophin hormone (ACTH) enhanced B release in incubating females. In males the adrenocortical response to stress was significantly attenuated during the parental phase compared with arrival at the breeding ground. In contrast to temperate passerines, there was no significant decrease in male territorial aggressive behaviour when B was experimentally elevated, suggesting a behavioural insensitivity to glucocorticoids. This mechanism is hypothesized to increase reproductive success by preventing interruptions to parental care during transient deleterious environmental perturbations, which are often experienced in the short Arctic breeding season. Modulation of the HPA axis in this species in relation to life-history stage, lifetime reproductive success and the polygynandrous mating system is discussed.
doi:10.1098/rspb.2003.2455
PMCID: PMC1691444  PMID: 12964988
13.  Familiarity with the experimenter influences the performance of Common ravens (Corvus corax) and Carrion crows (Corvus corone corone) in cognitive tasks☆ 
Behavioural Processes  2014;103(100):129-137.
Highlights
•We compared the results of corvids in experiments with familiar/unfamiliar humans.•We investigated behavioural reactions towards familiar and unfamiliar humans.•Corvids performed significantly better in experiments with familiar humans.•Corvids did not show more neophobia towards unfamiliar humans.•Hence, familiarity positively affected the experimental performance of corvids.
When humans and animals interact with one another over an extended time span they familiarise and may develop a relationship, which can exert an influence on both partners. For example, the behaviour of an animal in experiments may be affected by its relationship to the human experimenter. However, few studies have systematically examined the impact of human–animal relationships on experimental results. In the present study we investigated if familiarity with a human experimenter influences the performance of Common ravens (Corvus corax) and Carrion crows (Corvus corone corone) in interactive tasks. Birds were tested in two interactive cognitive tasks (exchange, object choice) by several experimenters representing different levels of familiarity (long and short-term). Our findings show that the birds participated more often in both tasks and were more successful in the exchange task when working with long-term experimenters than when working with short-term experimenters. Behavioural observations indicate that anxiety did not inhibit experimental performance but that the birds’ motivation to work differed between the two kinds of experimenters, familiar and less familiar. We conclude that human–animal relationships (i.e. familiarity) may affect the experimental performance of corvids in interactive cognitive tasks.
doi:10.1016/j.beproc.2013.11.013
PMCID: PMC4003535  PMID: 24333226
Carrion crow; Cognition; Common raven; Familiarity; Human–animal relationships
14.  Microarray-Based Transcriptomic Analysis of Differences between Long-Term Gregarious and Solitarious Desert Locusts 
PLoS ONE  2011;6(11):e28110.
Desert locusts (Schistocerca gregaria) show an extreme form of phenotypic plasticity and can transform between a cryptic solitarious phase and a swarming gregarious phase. The two phases differ extensively in behavior, morphology and physiology but very little is known about the molecular basis of these differences. We used our recently generated Expressed Sequence Tag (EST) database derived from S. gregaria central nervous system (CNS) to design oligonucleotide microarrays and compare the expression of thousands of genes in the CNS of long-term gregarious and solitarious adult desert locusts. This identified 214 differentially expressed genes, of which 40% have been annotated to date. These include genes encoding proteins that are associated with CNS development and modeling, sensory perception, stress response and resistance, and fundamental cellular processes. Our microarray analysis has identified genes whose altered expression may enable locusts of either phase to deal with the different challenges they face. Genes for heat shock proteins and proteins which confer protection from infection were upregulated in gregarious locusts, which may allow them to respond to acute physiological challenges. By contrast the longer-lived solitarious locusts appear to be more strongly protected from the slowly accumulating effects of ageing by an upregulation of genes related to anti-oxidant systems, detoxification and anabolic renewal. Gregarious locusts also had a greater abundance of transcripts for proteins involved in sensory processing and in nervous system development and plasticity. Gregarious locusts live in a more complex sensory environment than solitarious locusts and may require a greater turnover of proteins involved in sensory transduction, and possibly greater neuronal plasticity.
doi:10.1371/journal.pone.0028110
PMCID: PMC3223224  PMID: 22132225
15.  Color change as a potential behavioral strategy 
Hormones and behavior  2008;54(3):463-470.
Within species, color morphs may enhance camouflage, improve communication and/or confer reproductive advantage. However, in the male cichlid Astatotilapia burtoni, body color may also signal a behavioral strategy. A. burtoni live in a lek-like social system in Lake Tanganyika, Africa where bright blue or yellow territorial (T) males (together ~ 10–30% of the population) are reproductively capable and defend territories containing food with a spawning site. In contrast, nonterritorial (NT) males are smaller, cryptically colored, shoal with females and have regressed gonads. Importantly, males switch between these social states depending on their success in aggressive encounters. Yellow and blue morphs were thought to be adaptations to particular habitats, but they co-exist both in nature and in the laboratory. Importantly, individual males can switch colors so we asked whether color influences behavioral and hormonal profiles. When pairing territorial males with opposite colored fish, yellow males became dominant over blue males significantly more frequently. Moreover, yellow T males had significantly higher levels of 11-ketotosterone than blue T males while only blue NT males had higher levels of the stress hormone cortisol compared to the other groups. Thus color differences alone predict dominance status and hormone profiles in T males. Since T males can and do change color, this suggests that A. burtoni may use color as a flexible behavioral strategy.
doi:10.1016/j.yhbeh.2008.05.006
PMCID: PMC3019090  PMID: 18586245
16.  Physiological Stress Mediates the Honesty of Social Signals 
PLoS ONE  2009;4(3):e4983.
Background
Extravagant ornaments used as social signals evolved to advertise their bearers' quality. The Immunocompetence Handicap Hypothesis proposes that testosterone-dependent ornaments reliably signal health and parasite resistance; however, empirical studies have shown mixed support. Alternatively, immune function and parasite resistance may be indirectly or directly related to glucocorticoid stress hormones. We propose that an understanding of the interplay between the individual and its environment, particularly how they cope with stressors, is crucial for understanding the honesty of social signals.
Methodology/Principal Findings
We analyzed corticosterone deposited in growing feathers as an integrated measure of hypothalamic-pituitary-adrenal activity in a wild territorial bird, the red grouse Lagopus lagopus scoticus. We manipulated two key, interrelated components, parasites and testosterone, which influence both ornamentation and fitness. Birds were initially purged of parasites, and later challenged with parasites or not, while at the same time being given testosterone or control implants, using a factorial experimental design. At the treatment level, testosterone enhanced ornamentation, while parasites reduced it, but only in males not implanted with testosterone. Among individuals, the degree to which both parasites and testosterone had an effect was strongly dependent on the amount of corticosterone in the feather grown during the experiment. The more stressors birds had experienced (i.e., higher corticosterone), the more parasites developed, and the less testosterone enhanced ornamentation.
Conclusions/Significance
With this unique focus on the individual, and a novel, integrative, measure of response to stressors, we show that ornamentation is ultimately a product of the cumulative physiological response to environmental challenges. These findings lead toward a more realistic concept of honesty in signaling as well as a broader discussion of the concept of stress.
doi:10.1371/journal.pone.0004983
PMCID: PMC2656621  PMID: 19319197
17.  The effect of exogenous corticosterone on West Nile virus infection in Northern Cardinals (Cardinalis cardinalis) 
Veterinary Research  2012;43(1):34.
The relationship between stress and disease is thought to be unambiguous: chronic stress induces immunosuppression, which likely increases the risk of infection. However, this link has not been firmly established in wild animals, particularly whether stress hormones affect host responses to zoonotic pathogens, which can be transmitted to domesticated animal, wildlife and human populations. Due to the dynamic effects of stress hormones on immune functions, stress hormones may make hosts better or poorer amplifying hosts for a pathogen contingent on context and the host species evaluated. Using an important zoonotic pathogen, West Nile virus (WNV) and a competent host, the Northern Cardinal (Cardinalis cardinalis), we tested the effects of exogenous corticosterone on response to WNV infection. Corticosterone was administered at levels that individuals enduring chronic stressors (i.e., long-term inclement weather, food shortage, anthropogenic pollution) might experience in the wild. Corticosterone greatly impacted mortality: half of the corticosterone-implanted cardinals died between five - 11 days post-inoculation whereas only one of nine empty-implanted (control) birds died. No differences were found in viral titer between corticosterone- and empty-implanted birds. However, cardinals that survived infections had significantly higher average body temperatures during peak infection than individuals that died. In sum, this study indicates that elevated corticosterone could affect the survival of WNV-infected wild birds, suggesting that populations may be disproportionately at-risk to disease in stressful environments.
doi:10.1186/1297-9716-43-34
PMCID: PMC3372427  PMID: 22520572
18.  Spatial Distribution of Oak Mistletoe as It Relates to Habits of Oak Woodland Frugivores 
PLoS ONE  2014;9(11):e111947.
This study addresses the underlying spatial distribution of oak mistletoe, Phoradendron villosum, a hemi-parasitic plant that provides a continuous supply of berries for frugivorous birds overwintering the oak savanna habitat of California's outer coast range. As the winter community of birds consuming oak mistletoe varies from group-living territorial species to birds that roam in flocks, we asked if mistletoe volume was spatially autocorrelated at the scale of persistent territories or whether the patterns predicted by long-term territory use by western bluebirds are overcome by seed dispersal by more mobile bird species. The abundance of mistletoe was mapped on trees within a 700 ha study site in Carmel Valley, California. Spatial autocorrelation of mistletoe volume was analyzed using the variogram method and spatial distribution of oak mistletoe trees was analyzed using Ripley's K and O-ring statistics. On a separate set of 45 trees, mistletoe volume was highly correlated with the volume of female, fruit-bearing plants, indicating that overall mistletoe volume is a good predictor of fruit availability. Variogram analysis showed that mistletoe volume was spatially autocorrelated up to approximately 250 m, a distance consistent with persistent territoriality of western bluebirds and philopatry of sons, which often breed next door to their parents and are more likely to remain home when their parents have abundant mistletoe. Using Ripley's K and O-ring analyses, we showed that mistletoe trees were aggregated for distances up to 558 m, but for distances between 558 to 724 m the O-ring analysis deviated from Ripley's K in showing repulsion rather than aggregation. While trees with mistletoe were aggregated at larger distances, mistletoe was spatially correlated at a smaller distance, consistent with what is expected based on persistent group territoriality of western bluebirds in winter and the extreme philopatry of their sons.
doi:10.1371/journal.pone.0111947
PMCID: PMC4229145  PMID: 25389971
19.  Cryptic genetic variation and paraphyly in ravens. 
Widespread species that are morphologically uniform may be likely to harbour cryptic genetic variation. Common ravens (Corvus corax) have an extensive range covering nearly the entire Northern Hemisphere, but show little discrete phenotypic variation. We obtained tissue samples from throughout much of this range and collected mitochondrial sequence and nuclear microsatellite data. Our study revealed a deep genetic break between ravens from the western United States and ravens from throughout the rest of the world. These two groups, the 'California clade' and the 'Holarctic clade' are well supported and over 4% divergent in mitochondrial coding sequence. Microsatellites also reveal significant differentiation between these two groups. Ravens from Minnesota, Maine and Alaska are more similar to ravens from Asia and Europe than they are to ravens from California. The two clades come in contact over a huge area of the western United States, with mixtures of the two mitochondrial groups present in Washington, Idaho and California. In addition, the restricted range Chihuahuan raven (Corvus cryptoleucus) of the south-west United States and Mexico is genetically nested within the paraphyletic common raven. Our findings suggest that the common raven may have formerly consisted of two allopatric groups that may be in the process of remerging.
PMCID: PMC1690844  PMID: 11197122
20.  Ravens Intervene in Others’ Bonding Attempts 
Current Biology  2014;24(22):2733-2736.
Summary
The competition for power in a complex social world is hypothesized to be a driving force in the evolution of intelligence [1]. More specifically, power may be obtained not only by brute force but also by social strategies resembling human politics [2]. Most empirical evidence comes from primate studies that report unprovoked aggression by dominants to maintain power by spreading fear [3] and third-party interventions in conflicts [4–6]. Coalitionary support has also been described in other animals [7, 8] and is often linked to social bonding [9, 10]. As coalitions can lead to a gain in power [5, 11] and fitness benefits [12], individuals may try to prevent coalitionary support or indirectly prevent others from forming social bonds that might lead to coalitions. Although there is some empirical evidence that coalitionary support can be manipulated [13], little is known about the indirect strategy. We show here that wild ravens (Corvus corax) regularly intervene in affiliative interactions of others even though such interventions are potentially risky and without immediate benefits. Moreover, the identities of both interveners and intervened pairs are not randomly distributed. Ravens with existing ties initiate most interventions, and ravens that are creating new ties are most likely to be the targets of interventions. These patterns are consistent with the idea that interventions function to prevent others from forming alliances and consequently becoming future competitors. We thus show previously undescribed social maneuvers in the struggle for power. These maneuvers are likely to be of importance in other social species as well.
Highlights
•Ravens regularly engage in risky interventions in others’ affiliative encounters•Strongly bonded birds are more likely to intervene in affiliative interactions•Interventions almost exclusively target pairs that attempt to form new bonds•Ravens may prevent others from forming alliances and becoming future competitors
Animals can gain and maintain power by force, but also through support from social allies. Massen et al. show that ravens with established allies selectively intervene in bonding attempts of other ravens, suggesting that they try to prevent those from forming alliances. Such a “divide and rule” strategy has not been hitherto described for birds.
doi:10.1016/j.cub.2014.09.073
PMCID: PMC4245706  PMID: 25455033
21.  Social, state-dependent and environmental modulation of faecal corticosteroid levels in free-ranging female spotted hyenas. 
Little is known about to what extent the sensitivity of the hypothalamic-pituitary-adrenal (HPA) axis may be state dependent and vary in the same species between environments. Here we tested whether the faecal corticosteroid concentrations of matrilineal adult female spotted hyenas are influenced by social and reproductive status in adjacent ecosystems and whether they vary between periods with and without social stress. Females in the Serengeti National Park frequently become socially subordinate intruders in other hyena territories by undertaking long-distance foraging trips to migratory herds, whereas in the Ngorongoro Crater they usually forage inside their own small territories on resident prey. The faecal corticosteroid concentrations in Serengeti females were significantly higher than in Ngorongoro females. Energy expenditure by lactation is exceptionally high in spotted hyenas and this may be reflected in their corticosteroid levels. The faecal corticosteroid levels in both populations were higher in lactating than in non-lactating females. During periods of social stability, faecal corticosteroid concentrations increased in non-lactating females but not in lactating females as social status declined. Lactating Serengeti females had significantly higher faecal corticosteroid concentrations during periods with acute severe social stress than during periods without, indicating that the HPA axis is sensitive to social stimuli even in lactating females. So far few studies have used non-invasive monitoring methods for assessing social stress in freeranging animals. This study demonstrates for the first time, to the authors' knowledge, that corticosteroid concentrations may differ between periods with and without social stress for a free-ranging female mammal and that the modulating effect of social status may depend on reproductive status.
doi:10.1098/rspb.2001.1828
PMCID: PMC1088899  PMID: 11747563
22.  Parents are a Drag: Long-Lived Birds Share the Cost of Increased Foraging Effort with Their Offspring, but Males Pass on More of the Costs than Females 
PLoS ONE  2013;8(1):e54594.
Life history theory predicts that parents will balance benefits from investment in current offspring against benefits from future reproductive investments. Long-lived organisms are therefore less likely to increase parental effort when environmental conditions deteriorate. To investigate the effect of decreased foraging capacity on parental behaviour of long-lived monogamous seabirds, we experimentally increased energy costs for chick-rearing thick-billed murres (Uria lomvia). Handicapped birds had lighter chicks and lower provisioning rates, supporting the prediction that long-lived animals would pass some of the costs of impaired foraging ability on to their offspring. Nonetheless, handicapped birds spent less time underwater, had longer inter-dive surface intervals, had lower body mass, showed lower resighting probabilities in subsequent years and consumed fewer risky prey items. Corticosterone levels were similar between control and handicapped birds. Apparently, adults shared some of the costs of impaired foraging, but those costs were not measurable in all metrics. Handicapped males had higher plasma neutral lipid concentrations (higher energy mobilisation) and their chicks exhibited lower growth rates than handicapped females, suggesting different sex-specific investment strategies. Unlike other studies of auks, partners did not compensate for handicapping, despite good foraging conditions for unhandicapped birds. In conclusion, parental murres and their offspring shared the costs of experimentally increased foraging constraints, with females investing more than males.
doi:10.1371/journal.pone.0054594
PMCID: PMC3559872  PMID: 23382921
23.  Non-breeding feather concentrations of testosterone, corticosterone and cortisol are associated with subsequent survival in wild house sparrows 
Potential mechanistic mediators of Darwinian fitness, such as stress hormones or sex hormones, have been the focus of many studies. An inverse relationship between fitness and stress or sex hormone concentrations has been widely assumed, although empirical evidence is scarce. Feathers gradually accumulate hormones during their growth and provide a novel way to measure hormone concentrations integrated over time. Using liquid chromatography–tandem mass spectrometry, we measured testosterone, corticosterone and cortisol in the feathers of house sparrows (Passer domesticus) in a wild population which is the subject of a long-term study. Although corticosterone is considered the dominant avian glucocorticoid, we unambiguously identified cortisol in feathers. In addition, we found that feathers grown during the post-nuptial moult in autumn contained testosterone, corticosterone and cortisol levels that were significantly higher in birds that subsequently died over the following winter than in birds that survived. Thus, feather steroids are candidate prospective biomarkers to predict the future survival of individuals in the wild.
doi:10.1098/rspb.2011.2062
PMCID: PMC3282351  PMID: 22090380
corticosteroid; glucocorticoid; mass spectrometry; mortality; Passer domesticus; LC–MS/MS; stress
24.  Elevated Plasma Corticosterone Decreases Yolk Testosterone and Progesterone in Chickens: Linking Maternal Stress and Hormone-Mediated Maternal Effects 
PLoS ONE  2011;6(8):e23824.
Despite considerable research on hormone-mediated maternal effects in birds, the underlying physiology remains poorly understood. This study investigated a potential regulation mechanism for differential accumulation of gonadal hormones in bird eggs. Across vertebrates, glucocorticoids can suppress reproduction by downregulating gonadal hormones. Using the chicken as a model species, we therefore tested whether elevated levels of plasma corticosterone in female birds influence the production of gonadal steroids by the ovarian follicles and thus the amount of reproductive hormones in the egg yolk. Adult laying hens of two different strains (ISA brown and white Leghorn) were implanted subcutaneously with corticosterone pellets that elevated plasma corticosterone concentrations over a period of nine days. Steroid hormones were subsequently quantified in plasma and yolk. Corticosterone-implanted hens of both strains had lower plasma progesterone and testosterone levels and their yolks contained less progesterone and testosterone. The treatment also reduced egg and yolk mass. Plasma estrogen concentrations decreased in white Leghorns only whereas in both strains yolk estrogens were unaffected. Our results demonstrate for the first time that maternal plasma corticosterone levels influence reproductive hormone concentrations in the yolk. Maternal corticosterone could therefore mediate environmentally induced changes in yolk gonadal hormone concentrations. In addition, stressful situations experienced by the bird mother might affect the offspring via reduced amounts of reproductive hormones present in the egg as well as available nutrients for the embryo.
doi:10.1371/journal.pone.0023824
PMCID: PMC3160319  PMID: 21886826
25.  Differential reproductive responses to stress reveal the role of life-history strategies within a species 
Life-history strategies describe that ‘slow’- in contrast to ‘fast’-living species allocate resources cautiously towards reproduction to enhance survival. Recent evidence suggests that variation in strategies exists not only among species but also among populations of the same species. Here, we examined the effect of experimentally induced stress on resource allocation of breeding seabirds in two populations with contrasting life-history strategies: slow-living Pacific and fast-living Atlantic black-legged kittiwakes. We tested the hypothesis that reproductive responses in kittiwakes under stress reflect their life-history strategies. We predicted that in response to stress, Pacific kittiwakes reduce investment in reproduction compared with Atlantic kittiwakes. We exposed chick-rearing kittiwakes to a short-term (3-day) period of increased exogenous corticosterone (CORT), a hormone that is released during food shortages. We examined changes in baseline CORT levels, parental care and effects on offspring. We found that kittiwakes from the two populations invested differently in offspring when facing stress. In response to elevated CORT, Pacific kittiwakes reduced nest attendance and deserted offspring more readily than Atlantic kittiwakes. We observed lower chick growth, a higher stress response in offspring and lower reproductive success in response to CORT implantation in Pacific kittiwakes, whereas the opposite occurred in the Atlantic. Our findings support the hypothesis that life-history strategies predict short-term responses of individuals to stress within a species. We conclude that behaviour and physiology under stress are consistent with trade-off priorities as predicted by life-history theory. We encourage future studies to consider the pivotal role of life-history strategies when interpreting inter-population differences of animal responses to stressful environmental events.
doi:10.1098/rspb.2013.2090
PMCID: PMC3790493  PMID: 24089339
trade-offs; stress hormones; reproduction; populations; seabirds; life-history theory

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