Pigeon odor varies with experimental exposure to trace metal pollution.

Trace metals are chemical pollutants that have well-known noxious effects on wildlife and that are current major environmental issues in urban habitats. Previous studies have demonstrated their negative (e.g. lead) or positive (e.g. zinc) effects on body condition, immunity and reproductive success. Because of their effects on condition, trace metals are likely to influence the production of condition-dependent ornaments. The last decade has revealed that bird odors, like mammal odors, can convey information on individual quality and might be used as secondary sexual ornaments. Here, we used solid-phase microextraction headspace sampling with gas chromatography-mass spectrometry to investigate whether plumage scent varied with experimental supplementation in lead and/or zinc in feral pigeons. Zinc supplementation (alone or in combination with lead) changed the proportion of several volatiles, including an increase in the proportion of hydroxy-esters. The production of these esters, that most likely originate from preen gland secretions, may be costly and might thus be reduced by stress induced by zinc deficiency. Although lead is known to negatively impact pigeon condition, it did not statistically affect feather scent, despite most of the volatiles that increased with zinc exposure tended to be decreased in lead-supplemented pigeons. Further studies should evaluate the functions of plumage volatiles to predict how trace metals can impact bird fitness.

Do trace metals select for darker birds in urban areas? An experimental exposure to lead and zinc.

Trace metals from anthropogenic activities are involved in numerous health impairments and may therefore select for detoxification mechanisms or a higher tolerance. Melanin, responsible for the black and red colourations of teguments, plays a role in metal ion chelation and its synthesis is positively linked to immunity, antioxidant capacity and stress resistance due to pleiotropic effects. Therefore, we expected darker birds to (1) store higher amounts of metals in their feathers, (2) maintain lower metal concentrations in blood and (3) suffer less from metal exposure. We exposed feral pigeons (Columba livia) exhibiting various plumage darkness levels to low, but chronic, concentrations of zinc and/or lead, two of the most abundant metals in urban areas. First, we found negative and positive effects of lead and zinc, respectively, on birds' condition and reproductive parameters. Then, we observed positive relationships between plumage darkness and both zinc and lead concentrations in feathers. Interestingly, though darker adults did not maintain lower metal concentrations in blood and did not have higher fitness parameters, darker juveniles exhibited a higher survival rate than paler ones when exposed to lead. Our results show that melanin-based plumage colouration does modulate lead effects on birds' fitness parameters but that the relationship between metals, melanin, and fitness is more complex than expected and thus stress the need for more studies.

Sex-associated differences in trace metals concentrations in and on the plumage of a common urban bird species.

Urban areas encompass both favorable and stressful conditions linked with human activities and pollution. Pollutants remain of major ecological importance for synanthropic organisms living in the city. Plumage of urban birds harbour trace metals, which can result from external deposition or from internal accumulation. External and internal plumage concentrations likely differ between specific trace metals, and may further differ between males and females because of potential sex-linked differential urban use, physiology or behaviour. Here, we measured the concentrations in four trace metals (cadmium, copper, lead and zinc) in both unwashed and washed feathers of 49 male and 38 female feral pigeons (Columba livia) from Parisian agglomeration. We found that these concentrations indeed differed between unwashed and washed feathers, between males and females, and for some metals depended on the interaction between these factors. We discuss these results in the light of physiological and behavioural differences between males and females and of spatial repartition of the four trace metals in the city.

Mating modifies female life history in a haplodiploid spider mite.

Mating usually modifies females' resource allocation pattern, often as a result of conflicts between male and female partners. Can such a switch occur even in the absence of sexual conflicts? We addressed this issue in the haplodiploid spider mite Tetranychus urticae, whose biology and population structure considerably reduce conflicts between males and females over reproductive decisions. Comparing virgin and mated females, we tested the hypothesis that mated females modify their allocation pattern so as to maximize their probability of producing daughters. Mated females produced fewer but larger eggs, resulting in an overall similar reproductive effort but an increased probability of producing daughters, since in this species larger eggs are more likely to be fertilized and thus to become female. Moreover, mated females concentrated their reproduction early in life. Again, this might be a way to produce more daughters, since sperm is more abundant early in life. For virgins, spreading reproductive investment might be a way to save resources to extend life span, thus increasing their probability of encountering a sexual partner. Females with multiple opportunities for mating produced fewer eggs and a less female-biased sex ratio than once-mated females, raising the question of why multiple mating often occurs in this species.

Sex allocation in haplodiploids is mediated by egg size: evidence in the spider mite Tetranychus urticae Koch.

Haplodiploid species display extraordinary sex ratios. However, a differential investment in male and female offspring might also be achieved by a differential provisioning of eggs, as observed in birds and lizards. We investigated this hypothesis in the haplodiploid spider mite Tetranychus urticae, which displays highly female-biased sex ratios. We show that egg size significantly determines not only larval size, juvenile survival and adult size, but also fertilization probability, as in marine invertebrates with external fertilization, so that female (fertilized) eggs are significantly larger than male (unfertilized) eggs. Moreover, females with on average larger eggs before fertilization produce a more female-biased sex ratio afterwards. Egg size thus mediates sex-specific egg provisioning, sex and offspring sex ratio. Finally, sex-specific egg provisioning has another major consequence: male eggs produced by mated mothers are smaller than male eggs produced by virgins, and this size difference persists in adults. Virgin females might thus have a (male) fitness advantage over mated females.

Telomere erosion varies with sex and age at immune challenge but not with maternal antibodies in pigeons.

Conditions experienced early in life have profound impact on adult fitness, and telomere erosion could be a key mechanism in this process. In particular, early exposure to parasites is a frequent phenomenon in young vertebrates, which is associated with several short- and long-term costs such as telomere erosion. However, the timing of exposure to parasites during ontogeny and maternal antibodies can strongly modulate the costs of immunity, and could differentially affect telomere erosion. Here, we compared the effects of an early or late immune challenge on telomere erosion rate in male and female young feral pigeons (Columba livia) having received or not maternal antibodies. More specifically, we tested whether (i) early or late injections of antigens had different effects on nestling telomere erosion rate, (ii) whether this effect was different between male and female nestlings, and (iii) whether maternal antibodies could modulate telomere erosion rate. Our results show an interaction between sex and age at injection. Late-injected nestlings (injected at 14 days of age) had an accelerated erosion rate compared with the early-injected nestlings (injected at 3 days of age), and this effect was higher in females compared with the males. However, we did not find any effect of maternal antibodies on telomere erosion rate. These results suggest that the age at which an immune challenge occurs is important for telomere erosion and that sex-specific approaches are needed to better understand the short-term and long-term costs of parasite exposure in young vertebrates.

Oxidative stress responses in relationship to persistent organic pollutant levels in feathers and blood of two predatory bird species from Pakistan.

To date, knowledge of persistent organic pollutant (POP) mediated oxidative stress responses in avian species is rather limited. We therefore investigated whether exposure to polybrominated diphenyl ethers (PBDEs) and organochlorine pesticides (OCPs) in two predatory bird species, namely black kite (Milvus migrans) and spotted owlet (Athene brama), was associated to activities of antioxidant enzymes, such as glutathione peroxidase (GPx), glutathione S-transferase (GST), glutathione reductase (GR) and catalase (CAT), or expression of GPx and superoxide dismutase (SOD) genes. As part of this investigation, we evaluated whether feathers were suitable to reflect internal body burdens and their associated oxidative stress effects. p,p'-DDE was unanimously recorded with highest concentrations in feathers and blood of both species. In general, the non-significant associations reflect that feathers are not always a suitable indicator for internal body burdens of POPs, depending on the feather type and the age of the bird. The activity of GST and GR was significantly higher in spotted owlet whereas GPx and CAT was higher (albeit not significant) in spotted owlet and black kite respectively. In comparison, mRNA expression of GPx, SOD and Cu,ZnSOD was significantly higher in black kite. Regression analysis showed that the activity of GST and GR was significantly associated with p,p'-DDE in blood of spotted owlet. Similarly, activity of CAT and GR was significantly correlated with BDE-100 in feathers of spotted owlet. In comparison, mRNA expression of SOD was found significantly associated with ∑PBDEs in blood of spotted owlet as well as p,p'-DDE in feathers of black kite. Significant associations of various POPs with biological responses may suggest that POP exposure may be contributing to oxidative stress in the studied bird of prey species. This first investigation indicates the necessity for further research on cause-effect relationships between POP exposures and changes in general health of free ranging birds.

The first exposure assessment of legacy and unrestricted brominated flame retardants in predatory birds of Pakistan.

The exposure to legacy polybrominated diphenyl ethers (PBDEs), hexabromocyclododecanes (HBCDDs) and unrestricted 1,2-bis (2,4,6-tribromophenoxy) ethane (BTBPE), bis (2-ethylhexyl)-2,3,4,5-tetrabromophthalate (BEH-TEBP) and 2-ethylhexyl-2,3,4,5-tetrabromo-benzoate (EH-TBB) was examined in tail feathers of 76 birds belonging to ten predatory species inhabiting Pakistan. In addition, different feather types of six individuals of Black kite (Milvus migrans) were compared for their brominated flame retardant (BFR) levels. Black kite was found to be the most contaminated species with a median (minimum-maximum) tail feather concentration of 2.4 (0.70-7.5) ng g-1 dw for ∑PBDEs, 1.5 (0.5-8.1) ng g-1 dw for ∑HBCDDs and 0.10 ( 0.05 for both). Similarly, no significant concentration differences were observed among different feather types (all P > 0.05) suggesting their similar exposure. While variables such as species, trophic guild and δ15N values were evaluated as major predictors for BFR accumulation in the studied species, we predict that combined effects of just mentioned factors may govern the intra- and interspecific differences in BFR contamination profiles. We urge for further investigation of BFR exposure and potential toxicological effects in predatory birds from Asia with a more extensive sample size per species and location.

Temporal habitat variability and the maintenance of sex in host populations of the pea aphid.

The evolutionary maintenance of sex, despite competition from asexual reproduction, has long intrigued the evolutionary biologists owing to its numerous apparent short-term costs. In aphids, winter climate is expected to determine the maintenance of sexual lineages in the high latitude zones owing to their exclusive ability to produce frost-resistant eggs. However, diverse reproductive modes may coexist at a local scale where climatic influence is counteracted by microgeographical factors. In this study, we tested the influence of local habitat characteristics on regional coexistence of reproductive modes in the pea aphid, Acyrthosiphon pisum. In the laboratory, the induction of sexual morph production of many pea aphid genotypes from the local fields of annual (pea and faba bean) and perennial (alfalfa and red clover) crops in Western France indicated that A. pisum lineages from annual crops had a significantly higher investment in sexual reproduction than A. pisum lineages from the perennial hosts. We propose that temporal habitat variability exerts a selective pressure to maintain the sexual reproduction in A. pisum. The ecological and evolutionary consequences of the association between the mode of reproduction and the host population on gene flow restriction and on ecological specialization are discussed.

A review on current knowledge and future prospects of organohalogen contaminants (OHCs) in Asian birds.

The release of harmful chemicals in the Asian environment has recently increased dramatically due to rising industrial and agricultural activities. About 60% of the global human population is currently living on the Asian continent and may thus be exposed to a large range of different chemicals. Different classes of organohalogen chemicals have indeed been reported in various environmental compartments from Asia including humans and wildlife, but this issue has received less attention in birds. In this article, we reviewed the available literature on levels of legacy persistent organic pollutants (POPs) and various flame retardants (FRs) in Asian avifauna to analyze the existing pool of knowledge as well as to identify the gaps that should be addressed in future research. Furthermore, we discussed the variation in levels of organohalogens based on differences in regions, trophic level, dietary sources and migratory behaviors of species including distribution patterns in different tissues of birds. Although the mass of published literature is very low and even absent in many important regions of Asia, we deduced from the reported studies that levels of almost all classes of organohalogens (OHCs) including FRs were highest in East Asian countries such as Japan, China and South Korea, except for HCHs that were found at maximum levels in birds of South India. Concentrations (ng/g LW) of different OHCs in Asian birds ranged between

Use of feathers to assess polychlorinated biphenyl and organochlorine pesticide exposure in top predatory bird species of Pakistan.

Little is known about the levels of organochlorines (OCs) in predatory bird species from Asia or the factors governing their concentrations. This study is the first report on concentrations of polychlorinated biphenyls (PCBs) and organochlorine pesticides (OCPs) in predatory birds of Pakistan. The concentrations of PCBs and OCPs were investigated using tail feathers of ten different species of predatory birds. In addition, concentration differences among body, tail, primary and secondary feathers were investigated for six individuals of black kite (Milvus migrans). Ranges of concentrations were highest for dichlorodiphenyldichloroethylene (p,p'-DDE: 0.11-2163ngg(-1) dry wt.) followed by dichlorodiphenyltrichloroethane (p,p'-DDT: 0.36-345ngg(-1) dry wt.), hexachlorobenzene (HCB: 0.02-34ngg(-1) dry wt.), ∑PCBs (0.03-16ngg(-1) dry wt.) and trans-nonachlor (TN; 0.01-0.13ngg(-1) dry wt.). CB 118, 153, 138, and 180 along with p,p'-DDE were found as the most prevalent compounds. ∑PCBs and ∑DDTs were significantly different among species (both p<0.01) and omnivorous, scavengers, carnivorous and piscivorous trophic guilds (all p<0.03). Only ∑PCBs were significantly differentamong different families of birds (p<0.01). Values of stable isotopes (δ(13)C and δ(15)N) differed significantly (all p<0.01) among species, families, trophic guilds as well as terrestrial and aquatic habitat but not between nocturnal and diurnal predators (p=0.22 for δ(13)C; p=0.50 for δ(15)N). Concentrations of ∑PCBs, ∑DDTs and trans-nonachlor, but not HCB (p=0.86), were significantly different among different feather types (all p<0.01). Trophic and taxonomic affiliation as well as dietary carbon sources (δ(13)C) for species were identified as the variables best explaining the observed variation in exposure to the studied compounds. The significance of contributing factors responsible for OC contamination differences in predatory birds should be further elucidated in future studies.

Color differences among feral pigeons (Columba livia) are not attributable to sequence variation in the coding region of the melanocortin-1 receptor gene (MC1R).

BACKGROUND: Genetic variation at the melanocortin-1 receptor (MC1R) gene is correlated with melanin color variation in many birds. Feral pigeons (Columba livia) show two major melanin-based colorations: a red coloration due to pheomelanic pigment and a black coloration due to eumelanic pigment. Furthermore, within each color type, feral pigeons display continuous variation in the amount of melanin pigment present in the feathers, with individuals varying from pure white to a full dark melanic color. Coloration is highly heritable and it has been suggested that it is under natural or sexual selection, or both. Our objective was to investigate whether MC1R allelic variants are associated with plumage color in feral pigeons. FINDINGS: We sequenced 888 bp of the coding sequence of MC1R among pigeons varying both in the type, eumelanin or pheomelanin, and the amount of melanin in their feathers. We detected 10 non-synonymous substitutions and 2 synonymous substitution but none of them were associated with a plumage type. It remains possible that non-synonymous substitutions that influence coloration are present in the short MC1R fragment that we did not sequence but this seems unlikely because we analyzed the entire functionally important region of the gene. CONCLUSIONS: Our results show that color differences among feral pigeons are probably not attributable to amino acid variation at the MC1R locus. Therefore, variation in regulatory regions of MC1R or variation in other genes may be responsible for the color polymorphism of feral pigeons.

Contrasting levels of heavy metals in the feathers of urban pigeons from close habitats suggest limited movements at a restricted scale.

Despite restrictions in emissions, heavy metals may remain a major environmental issue due to their numerous sources and their persistence. Here, we assessed current levels of 4 metals (Copper, Cadmium, Lead, Zinc) in the feathers of 91 feral pigeons (Columba livia) from 7 sites in the urbanized region of Paris. Elements were detected in all pigeons, indicating that metals persist in urbanized areas. The ratio between metal concentrations in the feathers vs. in the environment calculated using data from other studies was 2-90 times higher for cadmium than for other metals, underlying its ecological importance. Concentrations in the feathers depended on locality, suggesting that pigeons remain in local habitats at this restricted scale, as expected from previous observations. Overall, our study suggests that urban feral pigeons may represent a good model system for metal biomonitoring.

Facultative symbiont infections affect aphid reproduction.

Some bacterial symbionts alter their hosts reproduction through various mechanisms that enhance their transmission in the host population. In addition to its obligatory symbiont Buchnera aphidicola, the pea aphid Acyrthosiphon pisum harbors several facultative symbionts influencing several aspects of host ecology. Aphids reproduce by cyclical parthenogenesis whereby clonal and sexual reproduction alternate within the annual life cycle. Many species, including the pea aphid, also show variation in their reproductive mode at the population level, with some lineages reproducing by cyclical parthenogenesis and others by permanent parthenogenesis. While the role of facultative symbionts has been well studied during the parthenogenetic phase of their aphid hosts, very little is known on their possible influence during the sexual phase. Here we investigated whether facultative symbionts modulate the capacity to produce sexual forms in various genetic backgrounds of the pea aphid with controlled symbiont composition and also in different aphid genotypes from natural populations with previously characterized infection status and reproductive mode. We found that most facultative symbionts exhibited detrimental effects on their hosts fitness under sex-inducing conditions in comparison with the reference lines. We also showed that the loss of sexual phase in permanently parthenogenetic lineages of A. pisum was not explained by facultative symbionts. Finally, we demonstrated that Spiroplasma infection annihilated the production of males in the host progeny by inducing a male-killing phenotype, an unexpected result for organisms such as aphids that reproduce primarily through clonal reproduction.

Strong biases in the transmission of sex chromosomes in the aphid Rhopalosiphum padi.

The typical life cycle of aphids involves several parthenogenetic generations followed by a single sexual one in autumn, i.e. cyclical parthenogenesis. Sexual females are genetically identical to their parthenogenetic mothers and carry two sex chromosomes (XX). Male production involves the elimination of one sex chromosome (to produce X0) that could give rise to genetic conflicts between X-chromosomes. In addition, deleterious recessive mutations could accumulate on sex chromosomes during the parthenogenetic phase and affect males differentially depending on the X-chromosome they inherit. Genetic conflicts and deleterious mutations thus may induce transmission bias that could be exaggerated in males. Here, the transmission of X-chromosomes has been studied in the laboratory in two cyclically parthenogenetic lineages of the bird cherry-oat aphid Rhopalosiphum padi . X-chromosome transmission was followed, using X-linked microsatellite loci, at male production in the two lineages and in their hybrids deriving from reciprocal crosses. Genetic analyses revealed non-Mendelian inheritance of X-chromosomes in both parental and hybrid lineages at different steps of male function. Putative mechanisms and evolutionary consequences of non-Mendelian transmission of X-chromosomes to males are discussed.