Adrien Frantz

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.

Ecological specialization correlates with genotypic differentiation in sympatric host‐populations of the pea aphid

The pea aphid, Acyrthosiphon pisum, encompasses distinct host races specialized on various Fabaceae species, but the extent of genetic divergence associated with ecological specialization varies greatly depending on plant and geographic origins of aphid populations. Here, we studied the genetic structure of French sympatric pea aphid populations collected on perennial (pea and faba bean) and annual (alfalfa and red clover) hosts using 14 microsatellite loci. Classical and Bayesian population genetics analyses consistently identified genetic clusters mostly related to plant origin: the pea/faba bean cluster was highly divergent from the red clover and the alfalfa ones, indicating they represent different stages along the continuum of genetic differentiation. Some genotypes were assigned to a cluster differing from the one expected from their plant origin while others exhibited intermediate genetic characteristics. These results suggest incomplete barriers to gene flow. However, this limited gene flow seems insufficient to prevent ecological specialization and genetic differentiation in sympatry.

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.

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.

The adaptive function of melanin-based plumage coloration to trace metals

Trace metals produced by anthropogenic activities are of major importance in urban areas and might constitute a new evolutionary force selecting for the ability to cope with their deleterious effects. Interestingly, melanin pigments are known to bind metal ions, thereby potentially sequestering them in inert body parts such as coat and feathers, and facilitating body detoxification. Thus, a more melanic plumage or coat coloration could bring a selective advantage for animals living in polluted areas. We tested this hypothesis by investigating the link between melanin-based coloration and zinc and lead concentrations in feathers of urban feral pigeons, both at capture time and after one year of captivity in standardized conditions. Results show that differently coloured pigeons had similar metal concentrations at capture time. Metal concentrations strongly decreased after one year in standardized conditions, and more melanic pigeons had higher concentrations of zinc (but not lead) in their feathers. This suggests that more melanic pigeons have a higher ability to store some metals in their feathers compared with their paler counterparts, which could explain their higher success in urbanized areas. Overall, this work suggests that trace metal pollution may exert new selective forces favouring more melanic phenotypes in polluted environments.

A potential role for parasites in the maintenance of color polymorphism in urban birds

Urbanization is a major challenge for biodiversity conservation, yet the evolutionary processes taking place in urbanized areas remain poorly known. Human activities in cities set new selective forces in motion which need to be investigated to predict the evolutionary responses of animal species living in urban areas. In this study, we investigated the role of urbanization and parasites in the maintenance of melanin-based color polymorphism in the feral pigeon Columba livia. Using a correlative approach, we tested whether differently colored genotypes displayed alternative phenotypic responses to urbanization, by comparing body condition, blood parasite prevalence and parasite load between colored morphs along an urbanization gradient. Body condition did not vary with urbanization, but paler individuals had a higher body condition than darker individuals. Moreover, paler morphs were less often parasitized than darker morphs in moderately urbanized habitats, but their parasite prevalence increased with urbanization. In contrast, darker morphs had similar parasite prevalence along the urbanization gradient. This suggests that paler morphs did better than darker morphs in moderately urbanized environments but were negatively affected by increasing urbanization, while darker morphs performed equally in all environments. Thus, differently colored individuals were distributed non-randomly across the urban habitat and suffered different parasite risk according to their location (a gene-by-environment interaction). This suggests that melanin-based coloration might reflect alternative strategies to cope with urbanization via different exposure or susceptibility to parasites. Spatial variability of parasite pressures linked with urbanization may, thus, play a central role in the maintenance of plumage color polymorphism in this urban 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.

Impact of urban environment and host phenotype on the epidemiology of Chlamydiaceae in feral pigeons (Columba livia)

Chlamydiaceae are obligate intracellular Gram-negative bacteria found all over the world and known to cause various forms of disease in animals and humans. Urban pigeons are known to be an important reservoir of Chlamydia psittaci, the agent of human psittacosis. In this study, we examined the influence of pigeon houses used to regulate pigeon populations and of melanin-based coloration on several epidemiological parameters of Chlamydiaceae in 708 urban pigeons in Paris. We also identified species and genotypes of Chlamydiaceae present in Parisian populations. First, our results revealed that pigeons roosting and breeding in pigeon houses were equally infected by Chlamydiaceae as those that did not. Second, we found that dark melanic pigeons excreted more Chlamydiaceae than pale melanic ones. Finally, species and strain diversities were very low: all samples were of C. psittaci genotype B. Nevertheless, two atypical Chlamydiaceae were identified based on 16S rRNA and ompA sequences. Our study thus highlights the importance of considering environmental and host phenotype when investigating the epidemiology of infectious diseases.

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 <LOD (limit of detection) to 14,000,000 for polychlorinated biphenyls (PCBs), <LOD to 790,000 for dichlorodiphenyltrichloroethane (DDTs), <LOD to 12,000 for hexachlorobenzene (HCB), <LOD to 29,000 for hexachlorocyclohexanes (HCHs), <LOD to 47,000 for chlordanes (CHLs) and <LOD to 4600 for total cyclodienes. Further, ranges (ng/g LW) of 1.1 to 150,000 for Co-PCBs; <LOD to 27 for polychlorinated dibenzo-p-dioxins (PCDDs); <LOD to 45 for polychlorinated dibenzofurans (PCDFs) and 0.02 to 73 for PCDD/DFs have been reported in Asian aves. Among emerging FRs, levels of total polybrominated diphenyl ethers (PBDEs), total dechlorane plus (DPs) [syn and anti DPs] and hexabromocyclododecane (HBCDs) oscillated between <LOD to 134,000, <LOD to 3820 [<0.1-920 and <0.1-2900], and <LOD to 11,800 ng/g LW, respectively. Corresponding ranges of novel brominated flame retardants (nBFRs) such as decabromodiphenyl ethane (DBDPE) and 1,2-bis(2,4,6-tribromophenoxy)ethane (BTBPE) were <LOD to 820 and <LOD to 89 ng/g LW. Other nBFRs such as tetrabromobisphenol-A (TBBPA) hexabromobenzene (HBB) and pentabromoethylbenzene (PBEB) in Asian avifauna have been reported in very few studies. Dependence of organohalogens on dietary sources and subsequent biomagnification in the food chain has been corroborated through δ(15)N and δ(13)C stable isotope proxies. In general, tissues with higher fat content accumulated more organohalogens and vice versa. Aspects such as maternal transfer of OHCs and temporal trends have rarely been discussed in reported literature from Asia. The mobility of birds, vicinity to sources and trans-boundary movement of pollutants were identified as key exposure routes and subsequent OHCs contamination in Asian birds. There is extreme scarcity of literature on organohalogen contamination in birds from Northern, South-eastern and west Asian countries where an industrial boom has been witnessed in the past few decades. Current scenarios suggest that levels of OHCs, particularly the FRs, are rising in birds of Asia and it would be wise to develop baseline information and to regulate the OHCs emission accordingly.

Host races of the pea aphid Acyrthosiphon pisum differ in male wing phenotypes.

The evolution of reproductive isolation without geographic isolation (sympatric speciation) has recently gained strong theoretical and empirical supports. It is now widely admitted that many host-specific phytophagous insect species have arisen through shifting and adapting to new plants. The pea aphid Acyrthosiphon pisum has received considerable attention in this context and is now considered as a probable case of incipient sympatric speciation through host specialization. In Europe, three host races have been described so far, one on annual plants (pea and broad bean) and two on perennial plants (red clover and alfalfa, respectively). These host races are genetically differentiated and exhibit strong ecological specialization affecting their preferences and performances on alternative plants. Here, we investigate whether other life-history traits of ecological importance are associated with host specialization in the species. In particular, because A. pisum shows a genetically determined male wing variation, we tested if its host races also differ in their proportion of winged/wingless male phenotypes. We used a large collection of pea aphid lineages sampled on pea, broad bean, red clover and alfalfa and analyzed their male production by placing them in conditions inducing the sexual phase in A. pisum. Striking differences in the frequency of male dispersal genotypes were found between host populations; aphids producing winged males were in high proportion among lineages from annual hosts, while those producing wingless males were in high proportion on perennial ones. The evolutionary maintenance and ecological consequences of this association between habitat specialization and male wing variation are discussed.