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1.  Foraging for carotenoids: do colorful male hihi target carotenoid-rich foods in the wild? 
Behavioral Ecology  2014;25(5):1048-1057.
Lay Summary
Birds that color their feathers with dietary carotenoid pigments are expected to seek out these pigments when they are molting. We show that molting male hihi, who express carotenoid-based plumage, seek out naturally occurring foods that are rich in carotenoid pigments. Female hihi, who do not express carotenoid-based plumage, do not seek out carotenoid-rich foods. This lends strength to the idea that carotenoid-based plumage reveals an individual’s foraging ability.
Dietary access to carotenoids is expected to determine the strength of carotenoid-based signal expression and potentially to maintain signal honesty. Species that display carotenoid-based yellow, orange, or red plumage are therefore expected to forage selectively for carotenoid-rich foods when they are depositing these pigments during molt, but whether they actually do so is unknown. We set out to address this in the hihi (Notiomystis cincta), a New Zealand passerine where males, but not females, display yellow carotenoid-based plumage. We measured circulating carotenoid concentrations in male and female hihi during breeding and molt, determined the nutritional content of common foods in the hihi diet, and conducted feeding observations of male and female hihi during molt. We found that although male and female hihi do not differ significantly in plasma carotenoid concentration, male hihi have a greater proportion of carotenoid-rich foods in their diet than do females. This is a consequence of a greater fruit and lower invertebrate intake than females and an avoidance of low-carotenoid content fruit. By combining behavioral observations with quantification of circulating carotenoids, we present evidence that colorful birds forage to maximize carotenoid intake, a conclusion we would not have drawn had we examined plasma carotenoids alone.
doi:10.1093/beheco/aru076
PMCID: PMC4160110  PMID: 25214753
carotenoids; foraging; hihi; nutrition; plumage.
2.  Female-specific colouration, carotenoids and reproductive investment in a dichromatic species, the upland goose Chloephaga picta leucoptera 
Behavioral Ecology and Sociobiology  2010;64(11):1779-1789.
Although studies on the evolution and function of female ornaments have become more numerous in the last years, the majority of these studies were carried out in cases where female ornaments were a smaller and duller version of the ornaments found in males. There are substantially fewer studies on species with female-specific ornaments. However, no study so far investigated the potential of female-specific colouration as a quality signal in birds with conventional sex roles. We studied female-specific ornamentation in a strongly sexually dichromatic species, the upland goose Chloephaga picta leucoptera, in two consecutive years. Male upland geese have white head and breast feathers and black legs, whereas females have reddish-brown head and breast feathers and conspicuous yellow-orange legs. We found that female-specific colouration in upland geese can reliably indicate different aspects of female phenotypic quality. Females with more orange coloured legs and more red-like head colours had higher clutch and egg volumes than females with a paler leg and head colouration, and a more reddish plumage colouration was related to a higher body condition. These relationships provide the theoretic possibility for males to assess female phenotypic quality on the basis of colouration. Furthermore, the females with a more orange-like tarsus colouration had higher plasma carotenoid levels. Both tarsus colouration and carotenoid concentrations of individual females were highly correlated across years, indicating that tarsus colour is a stable signal. Despite this correlation, small individual differences in plasma carotenoid concentrations between the two study years were related to differences in tarsus colouration. We thus show for the first time in a wild bird and under natural conditions that carotenoid-based integument colouration remains consistent between individuals in consecutive years and is also a dynamic trait reflecting individual changes in carotenoid levels. In this species, where pairs form life-long bonds, the honesty of the carotenoid-based integument colouration suggests that it may be a sexually selected female ornament that has evolved through male mate choice.
Electronic supplementary material
The online version of this article (doi:10.1007/s00265-010-0990-4) contains supplementary material, which is available to authorized users.
doi:10.1007/s00265-010-0990-4
PMCID: PMC2952766  PMID: 20976290
Upland goose; Chloephaga picta leucoptera; Female-specific colouration; Signalling; Carotenoids; Individual quality
3.  Does Foraging Behaviour Affect Female Mate Preferences and Pair Formation in Captive Zebra Finches? 
PLoS ONE  2010;5(12):e14340.
Background
Successful foraging is essential for survival and reproductive success. In many bird species, foraging is a learned behaviour. To cope with environmental change and survive periods in which regular foods are scarce, the ability to solve novel foraging problems by learning new foraging techniques can be crucial. Although females have been shown to prefer more efficient foragers, the effect of males' foraging techniques on female mate choice has never been studied. We tested whether females would prefer males showing the same learned foraging technique as they had been exposed to as juveniles, or whether females would prefer males that showed a complementary foraging technique.
Methodology/Principal Findings
We first trained juvenile male and female zebra finches (Taeniopygia guttata) to obtain a significant proportion of their food by one of two foraging techniques. We then tested whether females showed a preference for males with the same or the alternative technique. We found that neither a male's foraging technique nor his foraging performance affected the time females spent in his proximity in the mate-choice apparatus. We then released flocks of these finches into an aviary to investigate whether assortative pairing would be facilitated by birds taught the same technique exploiting the same habitat. Zebra finches trained as juveniles in a specific foraging technique maintained their foraging specialisation in the aviary as adults. However, pair formation and nest location were random with regard to foraging technique.
Conclusions/Significance
Our findings show that zebra finches can be successfully trained to be foraging specialists. However, the robust negative results of the conditions tested here suggest that learned foraging specializations do not affect mate choice or pair formation in our experimental context.
doi:10.1371/journal.pone.0014340
PMCID: PMC3002270  PMID: 21179514
4.  Carotenoid scarcity, synthetic pteridine pigments and the evolution of sexual coloration in guppies (Poecilia reticulata). 
Carotenoid-based sexual coloration is the classic example of an honest signal of mate quality. Animals cannot synthesize carotenoid pigments and ultimately depend on dietary sources. Thus, in carotenoid-poor environments, carotenoid coloration may be a direct indicator of foraging ability and an indirect indicator of health and vigour. Carotenoid coloration may also be affected, more directly, by parasites in some species. Carotenoids are not, however, the only conspicuous pigments available to animals. Pteridine pigments, with similar spectral properties, are displayed in the exoskeletons and wings of insects, the irides of birds and the skins of fishes, lizards and amphibians. Unlike carotenoids, pteridines are synthesized de novo by animals. We report that the orange spots that male guppies (Poecilia reticulata) display to females contain red pteridine pigments (drosopterins) in addition to carotenoids. We also examined the relationship between drosopterin production by males and carotenoid availability in the field. The results contrasted sharply with the hypothesis that males use drosopterins to compensate for carotenoid scarcity: males used more, not less, drosopterins in streams with higher carotenoid availability. The positive association between drosopterin use and carotenoid availability could reflect the costs of drosopterin synthesis or it could be a consequence of females preferring a particular pigment ratio or hue. Male guppies appear to use drosopterin pigments in a manner that dilutes, but does not eliminate, the indicator value of carotenoid coloration.
doi:10.1098/rspb.2001.1624
PMCID: PMC1088733  PMID: 11410150
5.  The Effects of Dietary Carotenoid Supplementation and Retinal Carotenoid Accumulation on Vision-Mediated Foraging in the House Finch 
PLoS ONE  2011;6(6):e21653.
Background
For many bird species, vision is the primary sensory modality used to locate and assess food items. The health and spectral sensitivities of the avian visual system are influenced by diet-derived carotenoid pigments that accumulate in the retina. Among wild House Finches (Carpodacus mexicanus), we have found that retinal carotenoid accumulation varies significantly among individuals and is related to dietary carotenoid intake. If diet-induced changes in retinal carotenoid accumulation alter spectral sensitivity, then they have the potential to affect visually mediated foraging performance.
Methodology/Principal Findings
In two experiments, we measured foraging performance of house finches with dietarily manipulated retinal carotenoid levels. We tested each bird's ability to extract visually contrasting food items from a matrix of inedible distracters under high-contrast (full) and dimmer low-contrast (red-filtered) lighting conditions. In experiment one, zeaxanthin-supplemented birds had significantly increased retinal carotenoid levels, but declined in foraging performance in the high-contrast condition relative to astaxanthin-supplemented birds that showed no change in retinal carotenoid accumulation. In experiments one and two combined, we found that retinal carotenoid concentrations predicted relative foraging performance in the low- vs. high-contrast light conditions in a curvilinear pattern. Performance was positively correlated with retinal carotenoid accumulation among birds with low to medium levels of accumulation (∼0.5–1.5 µg/retina), but declined among birds with very high levels (>2.0 µg/retina).
Conclusion/Significance
Our results suggest that carotenoid-mediated spectral filtering enhances color discrimination, but that this improvement is traded off against a reduction in sensitivity that can compromise visual discrimination. Thus, retinal carotenoid levels may be optimized to meet the visual demands of specific behavioral tasks and light environments.
doi:10.1371/journal.pone.0021653
PMCID: PMC3126843  PMID: 21747917
6.  Brighter yellow blue tits make better parents. 
Whether or not bird ornaments are a signal for direct (e.g. good parents) or indirect (e.g. good genes) benefits to prospective partners in sexual selection is controversial. Carotene coloration in Parus species is directly related to the ingestion of caterpillars, so that a brightly carotene-coloured tit may be signalling its ability to find caterpillars, a main high-quality food source for good fledgling development, and hence its parental abilities. If carotene-based plumage coloration is related to the good-parent hypothesis, we predict that yellow plumage brightness of tit fathers should be positively correlated to their investment in offspring provisioning. Here, we use cross-fostering experiments in blue tits (Parus caeruleus) to show that chick development (as measured by tarsus length) is related to yellowness of the foster father, but not to that of the genetic parents. Using these data, we were able to measure, for the first time to our knowledge, the separate contribution of genetic and environmental factors (i.e. parental plumage coloration) to chick development, and hence parental investment. Our data, which relate carotenoid coloration to models of good parents, and data from other authors, which relate ultraviolet coloration to good-genes models, stress that different kinds of coloration within an individual may provide different units of information to prospective females.
doi:10.1098/rspb.2001.1882
PMCID: PMC1690890  PMID: 11839194
7.  Bill Redness Is Positively Associated with Reproduction and Survival in Male and Female Zebra Finches 
PLoS ONE  2012;7(7):e40721.
Sexual traits can serve as honest indicators of phenotypic quality when they are costly. Brightly coloured yellow to red traits, which are pigmented by carotenoids, are relatively common in birds, and feature in sexual selection. Carotenoids have been linked to immune and antioxidant function, and the trade-off between ornamentation and these physiological functions provides a potential mechanism rendering carotenoid based signals costly. Mutual ornamentation is also common in birds and can be maintained by mutual mate choice for this ornament or by a correlated response in one sex to selection on the other sex. When selection pressures differ between the sexes this can cause intralocus sexual conflict. Sexually antagonistic selection pressures have been demonstrated for few sexual traits, and for carotenoid-dependent traits there is a single example: bill redness was found to be positively associated with survival and reproductive output in male zebra finches, but negatively so in females. We retested these associations in our captive zebra finch population without two possible limitations of this earlier study. Contrary to the earlier findings, we found no evidence for sexually antagonistic selection. In both sexes, individuals with redder bills showed higher survival. This association disappeared among the females with the reddest bills. Furthermore, females with redder bills achieved higher reproductive output. We conclude that bill redness of male and female zebra finches honestly signals phenotypic quality, and discuss the possible causes of the differences between our results and earlier findings.
doi:10.1371/journal.pone.0040721
PMCID: PMC3395645  PMID: 22808243
8.  Availability of non-carotenoid antioxidants affects the expression of a carotenoid-based sexual ornament 
Biology Letters  2007;3(4):353-356.
Carotenoids are responsible for much of the yellow, orange and red pigmentation in the animal kingdom, and the importance of such coloration as an honest signal of individual quality has received widespread attention. In particular, owing to the multiple roles of carotenoids as pigments, antioxidants and immunostimulants, carotenoid-based coloration has been suggested to advertise an individual's antioxidant or immune defence capacity. However, it has recently been argued that carotenoid-based signals may in fact be advertising the availability of different antioxidants, many of which (including various vitamins, antioxidant enzymes and minerals) are colourless and so would be uninformative as components of a visual signal, yet often have greater biological activity than carotenoids. We tested this hypothesis by feeding male sticklebacks (Gasterosteus aculeatus) a diet containing a fixed level of carotenoids and either low or high, but biologically realistic levels of the colourless antioxidant vitamins C and E. High-antioxidant diet males produced significantly more intensely coloured (but not larger) carotenoid-based regions of nuptial coloration and were preferred over size-matched males of the opposite diet treatment in mate-choice trials. Furthermore, there were positive correlations between an individual's somatic antioxidant activity and signal intensity. Our data suggest that carotenoid-based ornaments may honestly signal an individual's availability of non-carotenoid antioxidants, allowing females to make adaptive mate-choice decisions.
doi:10.1098/rsbl.2007.0072
PMCID: PMC2390655  PMID: 17472903
α-tocopherol; ascorbic acid; oxidative stress; sexual signalling
9.  Parental preference for red mouth of chicks in a songbird 
Parental preferences during feeding and care-giving may select for ornamental traits in young, such as bright coloration. For chicks of coots, there is experimental evidence for this idea. We examined the hypothesis that bright yellow, orange and red mouths of chicks of songbirds have been favoured by feeding preferences in parents. In a field experiment, the orange–yellow mouths of great tit nestlings were dyed brightly red, and the feeding response of parents recorded. In nest boxes with extra daylight through a window, experimental chicks were on average given twice as much food (biomass) as control chicks (sham dyed). In normal nest boxes, the tendency was similar, but not significant. Thus, at least in good light, great tit parents prefer to feed young with red mouths, a preference for colourfulness that helps explain the evolution of bright gapes in chicks of songbirds (passerine birds).
doi:10.1098/rspb.1997.0132
PMCID: PMC1688550
Feeding Preferences Songbirds Mouth Colour
10.  Carotenoid-based coloration in cichlid fishes 
Animal colors play important roles in communication, ecological interactions and speciation. Carotenoid pigments are responsible for many yellow, orange and red hues in animals. Whereas extensive knowledge on the proximate mechanisms underlying carotenoid coloration in birds has led to testable hypotheses on avian color evolution and signaling, much less is known about the expression of carotenoid coloration in fishes. Here, we promote cichlid fishes (Perciformes: Cichlidae) as a system in which to study the physiological and evolutionary significance of carotenoids. Cichlids include some of the best examples of adaptive radiation and color pattern diversification in vertebrates. In this paper, we examine fitness correlates of carotenoid pigmentation in cichlids and review hypotheses regarding the signal content of carotenoid-based ornaments. Carotenoid-based coloration is influenced by diet and body condition and is positively related to mating success and social dominance. Gaps in our knowledge are discussed in the last part of this review, particularly in the understanding of carotenoid metabolism pathways and the genetics of carotenoid coloration. We suggest that carotenoid metabolism and transport are important proximate mechanisms responsible for individual and population-differences in cichlid coloration that may ultimately contribute to diversification and speciation.
doi:10.1016/j.cbpa.2014.03.006
PMCID: PMC4003536  PMID: 24667558
Pigment; Trade-off; Antioxidant; Signal; Cichlidae
11.  Dietary carotenoids predict plumage coloration in wild house finches. 
Carotenoid pigments are a widespread source of ornamental coloration in vertebrates and expression of carotenoid-based colour displays has been shown to serve as an important criterion in female mate choice in birds and fishes. Unlike other integumentary pigments, carotenoids cannot be synthesized; they must be ingested. Carotenoid-based coloration is condition-dependent and has been shown to be affected by both parasites and nutritional condition. A controversial hypothesis is that the expression of carotenoid-based coloration in wild vertebrates is also affected by the amount and types of carotenoid pigments that are ingested. We tested this carotenoid-limitation hypothesis by sampling the gut contents of moulting house finches and comparing the concentration of carotenoid pigments in their gut contents with the colour of growing feathers. We found a positive association: males that ingested food with a higher concentration of carotenoid pigments grew brighter ornamental plumage. We also compared the concentration of carotenoids in the gut contents of males from two subspecies of house finches with small and large patches of carotenoid-based coloration. Consistent with the hypothesis that carotenoid access drives the evolution of carotenoid-based colour displays, males from the population with limited ornamentation had much lower concentrations of carotenoids in their gut contents than males from the population with extensive ornamentation. These observations support the idea that carotenoid intake plays a part in determining the plumage brightness of male house finches.
doi:10.1098/rspb.2002.1980
PMCID: PMC1691014  PMID: 12061954
12.  Carotenoid trade-off between parasitic resistance and sexual display: an experimental study in the blackbird (Turdus merula) 
Many parasites depress the expression of the carotenoid-based colour displays of their hosts, and it has been hypothesized that animals face a trade-off in carotenoid allocation between immune functions and ‘degree of ornamentation’. While numerous correlative studies suggest that parasite infection decreases the intensity of carotenoid-based colour displays, the existence of this trade-off has never been demonstrated experimentally in a host–parasite model. In this study, we used the blackbird (Turdus merula) and Isospora (an intestinal parasite) to assess whether this trade-off does indeed exist. Blackbirds were supplemented with carotenoids while simultaneously being exposed to parasites. Supplemented males circulated more carotenoids in the blood and developed more brightly coloured bills than unsupplemented males. In addition, supplementation slowed down the replication rate of parasites. Supplementation with carotenoids enabled infected birds to maintain their bill coloration, whereas birds that were infected but not supplemented showed reduced bill coloration. At the same time, infection slowed carotenoid assimilation in the blood. Overall, we demonstrated that bill colour reflects a bird's health, and that only males with a carotenoid-rich diet are capable of coping with costs associated with parasitic infection. Carotenoids are thus traded off between host physiological response to parasites and secondary sexual traits. Further investigations are required to determine the physiological mechanisms that govern this trade-off.
doi:10.1098/rspb.2007.1383
PMCID: PMC2596825  PMID: 18055388
bill colour; carotenoids; coccidia; experimental infection; trade-off
13.  Do carotenoids buffer testosterone-induced immunosuppression? An experimental test in a colourful songbird 
Biology Letters  2007;3(4):375-378.
Testosterone (T) is hypothesized to be an important honesty reinforcer of animal sexual signals. Owing to its immunosuppressive effects, only those individuals that can immunologically withstand high T levels can develop the most exaggerated traits. To date, few studies have isolated phenotypic or genotypic buffers that provide ‘high-quality’ animals with such an advantage. Dietary carotenoid pigments may in fact confer such a benefit because when in high supply carotenoids boost immunocompetence and coloration in animals like birds and fishes. We examined the experimental effect of T elevation on carotenoid and immune status in male and female zebra finches (Taeniopygia guttata) and found that T was immunostimulatory in a generalized cell-mediated challenge. We also detected a significant interaction between T treatment and the change in plasma carotenoids that occurred during the immune challenge; the relationship between blood carotenoid change and immunity was positive in controls and negative in T-implanted birds. This suggests that, while correlationally birds with high carotenoid stores were inherently better at mounting strong immune responses, experimentally administered T induced birds to deplete carotenoids for maximizing their health. Our findings highlight a nutrient-specific mechanism by which animals escape high immune costs of T elevation and thus can still elevate ornamentation.
doi:10.1098/rsbl.2007.0190
PMCID: PMC2390673  PMID: 17550877
carotenoid coloration; hormones; immunity; Taeniopygia guttata; zebra finch
14.  Better red than dead: carotenoid-based mouth coloration reveals infection in barn swallow nestlings. 
Nestling birds solicit food from their parents by displaying their open brightly coloured gapes. Carotenoids affect gape colour, but also play a central role in immunostimulation. Therefore, we hypothesize that, by differentially allocating resources to nestlings with more brightly coloured gapes, parents favour healthy offspring which are able to allocate carotenoids to gape coloration without compromising their immune defence. We demonstrated that, in the barn swallow Hirundo rustica, (i) parents differentially allocate food to nestlings with an experimentally brighter red gape, (ii) nestlings challenged with a novel antigen (sheep red blood cells, SRBCs) have less bright gape colour than their control siblings, (iii) nestlings challenged with SRBCs but also provided with the principal circulating carotenoid (lutein) have more brightly coloured red gapes than their challenged but unsupplemented siblings and (iv) the gape colour of nestlings challenged with SRBCs and provisioned with lutein exceeds that of siblings that were unchallenged. This suggests that parents may favour nestlings with superior health by preferentially feeding offspring with the brightest gapes.
PMCID: PMC1690492  PMID: 10670953
15.  Immune Activation Reduces Sperm Quality in the Great Tit 
PLoS ONE  2011;6(7):e22221.
Mounting an immune response against pathogens incurs costs to organisms by its effects on important life-history traits, such as reproductive investment and survival. As shown recently, immune activation produces large amounts of reactive species and is suggested to induce oxidative stress. Sperm are highly susceptible to oxidative stress, which can negatively impact sperm function and ultimately male fertilizing efficiency. Here we address the question as to whether mounting an immune response affects sperm quality through the damaging effects of oxidative stress. It has been demonstrated recently in birds that carotenoid-based ornaments can be reliable signals of a male's ability to protect sperm from oxidative damage. In a full-factorial design, we immune-challenged great tit males while simultaneously increasing their vitamin E availability, and assessed the effect on sperm quality and oxidative damage. We conducted this experiment in a natural population and tested the males' response to the experimental treatment in relation to their carotenoid-based breast coloration, a condition-dependent trait. Immune activation induced a steeper decline in sperm swimming velocity, thus highlighting the potential costs of an induced immune response on sperm competitive ability and fertilizing efficiency. We found sperm oxidative damage to be negatively correlated with sperm swimming velocity. However, blood resistance to a free-radical attack (a measure of somatic antioxidant capacity) as well as plasma and sperm levels of oxidative damage (lipid peroxidation) remained unaffected, thus suggesting that the observed effect did not arise through oxidative stress. Towards the end of their breeding cycle, swimming velocity of sperm of more intensely colored males was higher, which has important implications for the evolution of mate choice and multiple mating in females because females may accrue both direct and indirect benefits by mating with males having better quality sperm.
doi:10.1371/journal.pone.0022221
PMCID: PMC3134482  PMID: 21765955
16.  Maternal investment of female mallards is influenced by male carotenoid-based coloration 
The differential allocation hypothesis predicts that females modify their investment in a breeding attempt according to its reproductive value. One prediction of this hypothesis is that females will increase reproductive investment when mated to high-quality males. In birds, it was shown that females can modulate pre-hatch reproductive investment by manipulating egg and clutch sizes and/or the concentrations of egg internal compounds according to paternal attractiveness. However, the differential allocation of immune factors has seldom been considered, particularly with an experimental approach. The carotenoid-based ornaments can function as reliable signals of quality, indicating better immunity or ability to resist parasites. Thus, numerous studies show that females use the expression of carotenoid-based colour when choosing mates; but the influence of this paternal coloration on maternal investment decisions has seldom been considered and has only been experimentally studied with artificial manipulation of male coloration. Here, we used dietary carotenoid provisioning to manipulate male mallard (Anas platyrhynchos) bill coloration, a sexually selected trait, and followed female investment. We show that an increase of male bill coloration positively influenced egg mass and albumen lysozyme concentration. By contrast, yolk carotenoid concentration was not affected by paternal ornamentation. Maternal decisions highlighted in this study may influence chick survival and compel males to maintain carotenoid-based coloration from the mate-choice period until egg-laying has been finished.
doi:10.1098/rspb.2010.1115
PMCID: PMC3030838  PMID: 20843851
maternal investment; yolk carotenoid concentration; egg mass; albumen lysozyme concentration; paternal ornamentation; carotenoid-based coloration
17.  The interplay between gonadal steroids and immune defence in affecting a carotenoid-dependent trait 
Behavioral Ecology and Sociobiology  2011;65(10):2007-2019.
The hypothesis that sexual ornaments are honest signals of quality because their expression is dependent on hormones with immune-depressive effects has received ambiguous support. The hypothesis might be correct for those signals that are carotenoid-dependent because the required carotenoid deposition in the signal, stimulated by testosterone, might lower the carotenoid-dependent immune defence of the organism. Two pathways underlying this androgen-dependent honest signaling have been suggested. Firstly, androgens that are needed for ornament expression may suppress immune defence, a cost that only high-quality animals can afford. Alternatively, immune activation may downregulate the production of androgens in low-quality individuals. Which of these alternatives is correct, and to what extent these effects are mediated by the different metabolites of androgens, remain open questions. To provide answers to these questions, we manipulated the levels of testosterone (T), 5α-dihydrotestosterone (DHT), and 17-β-estradiol (E2) in diamond doves Geopelia cuneata, a species in which both sexes exhibit a carotenoid-dependent, androgen-regulated red–orange periorbital ring of bare skin. On the first day of the experiment (day 0), we inserted steroid-releasing implants into groups of birds and on day 14, we subjected half of the birds to an immunological challenge by immunizing them with sheep red blood cells (SRBC). In females, but not in males, androgen but not estradiol treatments reduced antibody production to SRBC. In addition, the immunological challenge reduced redness and size of the trait as well as androgens levels in both sexes and in all treatments. This indicates that an immunological challenge can lower circulating T at the cost of the trait expression. These findings are in accordance with both pathways postulated in the immunocompetence-handicap hypothesis, but do not entirely support the idea that the immunosuppressive effect of androgens yields honest signaling since both T and DHT were not immunosuppressive in males, for which sexual signaling is supposed to be especially important.
doi:10.1007/s00265-011-1210-6
PMCID: PMC3172405  PMID: 21957328
ICHH; Ornament; Color; Testosterone; Estradiol; Dihydrotestosterone; SRBC; Humoral immunity
18.  Differential Effects of Early- and Late-Life Access to Carotenoids on Adult Immune Function and Ornamentation in Mallard Ducks (Anas platyrhynchos) 
PLoS ONE  2012;7(5):e38043.
Environmental conditions early in life can affect an organism’s phenotype at adulthood, which may be tuned to perform optimally in conditions that mimic those experienced during development (Environmental Matching hypothesis), or may be generally superior when conditions during development were of higher quality (Silver Spoon hypothesis). Here, we tested these hypotheses by examining how diet during development interacted with diet during adulthood to affect adult sexually selected ornamentation and immune function in male mallard ducks (Anas platyrhynchos). Mallards have yellow, carotenoid-pigmented beaks that are used in mate choice, and the degree of beak coloration has been linked to adult immune function. Using a 2×2 factorial experimental design, we reared mallards on diets containing either low or high levels of carotenoids (nutrients that cannot be synthesized de novo) throughout the period of growth, and then provided adults with one of these two diets while simultaneously quantifying beak coloration and response to a variety of immune challenges. We found that both developmental and adult carotenoid supplementation increased circulating carotenoid levels during dietary treatment, but that birds that received low-carotenoid diets during development maintained relatively higher circulating carotenoid levels during an adult immune challenge. Individuals that received low levels of carotenoids during development had larger phytohemagglutinin (PHA)-induced cutaneous immune responses at adulthood; however, dietary treatment during development and adulthood did not affect antibody response to a novel antigen, nitric oxide production, natural antibody levels, hemolytic capacity of the plasma, or beak coloration. However, beak coloration prior to immune challenges positively predicted PHA response, and strong PHA responses were correlated with losses in carotenoid-pigmented coloration. In sum, we did not find consistent support for either the Environmental Matching or Silver Spoon hypotheses. We then describe a new hypothesis that should be tested in future studies examining developmental plasticity.
doi:10.1371/journal.pone.0038043
PMCID: PMC3364205  PMID: 22666443
19.  Mate choice for a male carotenoid-based ornament is linked to female dietary carotenoid intake and accumulation 
Background
The coevolution of male traits and female mate preferences has led to the elaboration and diversification of sexually selected traits; however the mechanisms that mediate trait-preference coevolution are largely unknown. Carotenoid acquisition and accumulation are key determinants of the expression of male sexually selected carotenoid-based coloration and a primary mechanism maintaining the honest information content of these signals. Carotenoids also influence female health and reproduction in ways that may alter the costs and benefits of mate choice behaviours and thus provide a potential biochemical link between the expression of male traits and female preferences. To test this hypothesis, we manipulated the dietary carotenoid levels of captive female house finches (Carpodacus mexicanus) and assessed their mate choice behavior in response to color-manipulated male finches.
Results
Females preferred to associate with red males, but carotenoid supplementation did not influence the direction or strength of this preference. Females receiving a low-carotenoid diet were less responsive to males in general, and discrimination among the colorful males was positively linked to female plasma carotenoid levels at the beginning of the study when the diet of all birds was carotenoid-limited.
Conclusions
Although female preference for red males was not influenced by carotenoid intake, changes in mating responsiveness and discrimination linked to female carotenoid status may alter how this preference is translated into choice. The reddest males, with the most carotenoid rich plumage, tend to pair early in the breeding season. If carotenoid-related variations in female choice behaviour shift the timing of pairing, then they have the potential to promote assortative mating by carotenoid status and drive the evolution of carotenoid-based male plumage coloration.
doi:10.1186/1471-2148-12-3
PMCID: PMC3315416  PMID: 22233462
20.  Measuring Motivation for Appetitive Behaviour: Food-Restricted Broiler Breeder Chickens Cross a Water Barrier to Forage in an Area of Wood Shavings without Food 
PLoS ONE  2014;9(7):e102322.
Broiler breeders (parents of meat chickens) are selected for fast growth and become obese if fed ad libitum. To avoid this and maintain good health and reproductive ability, they are feed restricted to about 1/3 of what they would eat ad libitum. As a result, they experience chronic hunger and exhibit abnormal behaviour patterns that may indicate stress and frustration. One approach to measuring hunger is to observe how much birds will work, such as pecking a key, for access to more or different types of food. However, the sight, smell, and feedback from consumption of the feed reward changes the context and may artificially raise feeding motivation. To avoid this, we tested broiler breeders in an apparatus in which they could work for access to a wooden platform covered in wood shavings by crossing a water runway which increased in length and depth in 8 successive tests. In the wood shavings area, they could perform exploratory and foraging behaviour (the appetitive phase of feeding) but were never rewarded with feed. Sixty birds were divided into three feed quantity treatments: commercial restriction (R), and twice (2R) or three times (3R) this amount. Overall, birds fed R worked harder to reach the wood shavings area (reached it in a larger number of tests) than 2R and 3R birds (P<0.001). More restricted birds took less time to reach the area (P<0.001, R<2R<3R) and spent more time foraging while there (P<0.001, R>2R>3R). This indicates that restricted-fed birds were hungry and willing to work for the opportunity to forage even though food was never provided, suggesting that their motivation to perform the appetitive component of feeding behaviour (foraging/food searching) was sufficient to sustain their response. Thus food restriction in broiler breeders is a welfare concern. However these methods could be used to test alternative feeding regimes to attempt to find ways of alleviating hunger while still maintaining healthy growth and reproduction in these birds.
doi:10.1371/journal.pone.0102322
PMCID: PMC4113302  PMID: 25068283
21.  Free Radical Exposure Creates Paler Carotenoid-Based Ornaments: A Possible Interaction in the Expression of Black and Red Traits 
PLoS ONE  2011;6(4):e19403.
Oxidative stress could be a key selective force shaping the expression of colored traits produced by the primary animal pigments in integuments: carotenoids and melanins. However, the impact of oxidative stress on melanic ornaments has only recently been explored, whereas its role in the expression of carotenoid-based traits is not fully understood. An interesting study case is that of those animal species simultaneously expressing both kinds of ornaments, such as the red-legged partridge (Alectoris rufa). In this bird, individuals exposed to an exogenous source of free radicals (diquat) during their development produced larger eumelanin-based (black) plumage traits than controls. Here, we show that the same red-legged partridges exposed to diquat simultaneously developed paler carotenoid-based ornaments (red beak and eye rings), and carried lower circulating carotenoid levels as well as lower levels of some lipids involved in carotenoid transport in the bloodstream (i.e., cholesterol). Moreover, partridges treated with a hormone that stimulates eumelanin production (i.e., alpha-melanocyte-stimulating hormone) also increased blood carotenoid levels, but this effect was not mirrored in the expression of carotenoid-based traits. The redness of carotenoid-based ornaments and the size of a conspicuous eumelanic trait (the black bib) were negatively correlated in control birds, suggesting a physiological trade-off during development. These findings contradict recent studies questioning the sensitivity of carotenoids to oxidative stress. Nonetheless, the impact of free radicals on plasma carotenoids seems to be partially mediated by changes in cholesterol metabolism, and not by direct carotenoid destruction/consumption. The results highlight the capacity of oxidative stress to create multiple phenotypes during development through differential effects on carotenoids and melanins, raising questions about evolutionary constraints involved in the production of multiple ornaments by the same organism.
doi:10.1371/journal.pone.0019403
PMCID: PMC3083443  PMID: 21556328
22.  Melanin-based plumage coloration in the house finch is unaffected by coccidial infection 
For most species of birds, ornamental plumage coloration may result from two types of pigments: carotenoids and melanins. Despite the fact that melanin pigments can be synthesized by birds from basic, amino acid precursors, while carotenoids cannot be synthesized by birds and must be ingested, melanin-based plumage coloration and carotenoid-based plumage coloration have often been treated as a single trait in investigations of the function and evolution of plumage coloration. Expression of carotenoid-based coloration is known to be dependent on condition, while the effects of individual condition have not been well-tested for expression of melanin-based coloration. In this study, we experimentally tested the effect of coccidial infection of the intestinal tract of male house finches during moult on expression of melanin-based plumage coloration. Coccidial infection had a significant negative effect on carotenoid-based coloration, but it had no significant effect on melanin-based feather coloration. Unlike carotenoid-based coloration, melanin-based coloration may be cheap to produce, and honesty of melanin-based coloration my require social mediation.
doi:10.1098/rspb.1998.0405
PMCID: PMC1689168
23.  Novel methoxy-carotenoids from the burgundy-colored plumage of the Pompadour Cotinga Xipholena punicea 
Archives of biochemistry and biophysics  2010;504(1):10.1016/j.abb.2010.08.006.
Recent advances in the fields of chromatography, mass spectrometry, and chemical analysis have greatly improved the efficiency with which carotenoids can be extracted and analyzed from avian plumage. Prior to these technological developments, Brush (1968) [1] concluded that the burgundy-colored plumage of the male pompadour Cotinga Xipholena punicea is produced by a combination of blue structural color and red carotenoids, including astaxanthin, canthaxanthin, isozeaxanthin, and a fourth unidentified, polar carotenoid. However, X. punicea does not in fact exhibit any structural coloration. This work aims to elucidate the carotenoid pigments of the burgundy color of X. punicea plumage using advanced analytical methodology. Feathers were collected from two burgundy male specimens and from a third aberrant orange-colored specimen. Pigments were extracted using a previously published technique (McGraw et al. (2005) [2]), separated by high-performance liquid chromatography (HPLC), and analyzed by UV/Vis absorption spectroscopy, chemical analysis, mass spectrometry, nuclear magnetic resonance (NMR), and comparison with direct synthetic products. Our investigation revealed the presence of eight ketocarotenoids, including astaxanthin and canthaxanthin as reported previously by Brush (1968) [1]. Six of the ketocarotenoids contained methoxyl groups, which is rare for naturally-occurring carotenoids and a novel finding in birds. Interestingly, the carotenoid composition was the same in both the burgundy and orange feathers, indicating that feather coloration in X. punicea is determined not only by the presence of carotenoids, but also by interactions between the bound carotenoid pigments and their protein environment in the barb rami and barbules. This paper presents the first evidence of metabolically-derived methoxy-carotenoids in birds.
doi:10.1016/j.abb.2010.08.006
PMCID: PMC3887449  PMID: 20709013
Bird pigmentation; Carotenoid identification; Xanthophyll metabolism
24.  Is male rhesus macaque red color ornamentation attractive to females? 
Behavioral ecology and sociobiology  2014;68(7):1215-1224.
Male sexually-selected traits can evolve through different mechanisms: conspicuous and colorful ornaments usually evolve through inter-sexual selection, while weapons usually evolve through intra-sexual selection. Male ornaments are rare among mammals in comparison to birds, leading to the notion that female mate choice generally plays little role in trait evolution in this taxon. Supporting this view, when ornaments are present in mammals they typically indicate social status and are products of male-male competition. This general mammalian pattern, however, may not apply to rhesus macaques (Macaca mulatta). Males of this species display conspicuous skin coloration, but this expression is not correlated to dominance rank, and is therefore unlikely to have evolved due to male-male competition. Here, we investigate whether male color expression influences female proceptivity towards males in the Cayo Santiago free-ranging rhesus macaque population. We collected face images of 24 adult males varying in dominance rank and age at the peak of the mating season, and modeled these to rhesus macaque visual perception. We also recorded female socio-sexual behaviors towards these males. Results show that dark red males received more sexual solicitations, by more females, than pale pink ones. Together with previous results, our study suggests that male color ornaments are more likely to be a product of inter- rather than intra-sexual selection. This may especially be the case in rhesus macaques due to the particular characteristics of male-male competition in this species.
doi:10.1007/s00265-014-1732-9
PMCID: PMC4167843  PMID: 25246728
Ornaments; sexual selection; female mate choice; sexual skin; color; anthropoid primates
25.  Blood parasite infection differentially relates to carotenoid-based plumage and bill color in the American goldfinch 
Ecology and Evolution  2014;4(16):3210-3217.
Male and female American goldfinches (Spinus tristis) express condition-dependent carotenoid-based plumage and bill coloration. Plumage color is relatively static, as pigments incorporated into feathers during the spring molt cannot be mobilized thereafter. In contrast, bill color is dynamic, reflecting changes in condition over short time periods. Previous studies have shown that male and female ornaments, though similar in expression, are differentially related to measures of immunocompetence, suggesting that the relationship between ornamentation and parasite infection may differ between the sexes. In this study, we evaluate the relationship between condition-dependent ornamentation (plumage and bill color) and blood parasite infection in male and female American goldfinches. We captured goldfinches after completion of the pre-alternate molt and prior to the onset of nesting and assessed prevalence of Trypanosoma parasites via blood smears. Plumage color strongly predicted trypanosome infection: Birds with more colorful plumage were less likely to present infections. In contrast, we detected no relationship between infection and bill color, which in other studies has been shown to dynamically reflect current condition. Sex did not affect the relationship between infection status and either ornament. Together, these results suggest that physiological pathways linking carotenoid ornamentation and infection may vary even within a single species.
doi:10.1002/ece3.1164
PMCID: PMC4222208  PMID: 25473474
American goldfinch; bill color; blood parasites; carotenoid-based ornamentation; plumage color; Spinus tristis; Trypanosoma

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