Malaria infections reinforce competitive asymmetry between two Ficedula flycatchers in a recent contact zone

Parasites may influence the outcome of interspecific competition between closely related host species through lower parasite virulence in the host with which they share the longer evolutionary history. We tested this idea by comparing the prevalence of avian malaria (Haemosporidia) lineages and their association with survival in pied and collared flycatchers (Ficedula hypoleuca and F. albicollis) breeding in a recent contact zone on the Swedish island of Öland. A nested PCR protocol amplifying haemosporidian fragments of mtDNA was used to screen the presence of malaria lineages in 1048 blood samples collected during 6 years. Competitively inferior pied flycatchers had a higher prevalence of blood parasites, including the lineages that were shared between the two flycatcher species. Multistate mark–recapture models revealed a lower survival of infected versus uninfected female pied flycatchers, while no such effects were detected in male pied flycatchers or in collared flycatchers of either sex. Our results show that a comparatively new host, the collared flycatcher, appears to be less susceptible to a local northern European malarial lineage where the collared flycatchers have recently expanded their distribution. Pied flycatchers experience strong reproductive interference from collared flycatchers, and the additional impact of species‐specific blood parasite effects adds to this competitive exclusion. These results support the idea that parasites can strongly influence the outcome of interspecific competition between closely related host species, but that the invading species need not necessarily be more susceptible to local parasites.     

Inferring the ecology of willow warblers during their winter moult by sequential stable isotope analyses of remiges

We present a comparison of feather stable isotope (δ¹³C, δ¹⁵N) patterns representing the habitat and diet conditions for two subspecies of willow warblers Phylloscopus trochilus that breed in parapatry, but winter in different regions of sub‐Saharan Africa. Previous analyses have shown that on average winter moulted innermost primaries (P1) show subspecific differences in δ¹⁵N values, although individuals show substantial variation for both δ¹³C and δ¹⁵N within the subspecies. We examined whether corresponding variation in the timing of the winter moult, as reflected by consistent intra‐wing correlations for individual's δ¹³C and δ¹⁵N values, could explain some of the previously observed isotopic variation. Further, differential subspecific adaptations to winter precipitation patterns across Africa might result in a variable degree of site fidelity or itinerancy during moult. We found no consistent trend in isotopic values from innermost to outermost primaries, thus inter‐individual variation in the timing of moult does not explain the subspecific isotopic variation for P1. Patterns in wing feather δ¹³C and δ¹⁵N values indicated that 41% of the individuals from both subspecies shifted their diet or habitats during winter moult. Importantly, despite well‐documented itinerancy in willow warblers during the winter, 59% of the individuals had feather isotope values consistent with stable use of habitats or diets during winter moult. Repeatability analyses suggest that individuals of both subspecies initiate moult in similar habitats from year‐to‐year while feeding on isotopically similar diets.     

Migration distance does not predict blood parasitism in a migratory songbird

Migration can influence host–parasite dynamics in animals by increasing exposure to parasites, by reducing the energy available for immune defense, or by culling of infected individuals. These mechanisms have been demonstrated in several comparative analyses; however, few studies have investigated whether conspecific variation in migration distance may also be related to infection risk. Here, we ask whether autumn migration distance, inferred from stable hydrogen isotope analysis of summer‐grown feathers (δ²H_f) in Europe, correlates with blood parasite prevalence and intensity of infection for willow warblers (Phylloscopus trochilus) wintering in Zambia. We also investigated whether infection was correlated with individual condition (assessed via corticosterone, scaled mass index, and feather quality). We found that 43% of birds were infected with Haemoproteus palloris (lineage WW1). Using generalized linear models, we found no relationship between migration distance and either Haemoproteus infection prevalence or intensity. There was spatial variation in breeding ground origins of infected versus noninfected birds, with infected birds originating from more northern sites than noninfected birds, but this difference translated into only slightly longer estimated migration distances (~214 km) for infected birds. We found no relationship between body condition indices and Haemoproteus infection prevalence or intensity. Our results do not support any of the proposed mechanisms for migration effects on host–parasite dynamics and cautiously suggest that other factors may be more important for determining individual susceptibility to disease in migratory bird species.     

Good genes, oxidative stress and condition-dependent sexual signals

The immune and the detoxication systems of animals are characterized by allelic polymorphisms, which underlie individual differences in ability to combat assaults from pathogens and toxic compounds. Previous studies have shown that females may improve offspring survival by selecting mates on the basis of sexual ornaments and signals that honestly reveal health. In many cases the expression of these ornaments appears to be particularly sensitive to oxidative stress. Activated immune and detoxication systems often generate oxidative stress by an extensive production of reactive metabolites and free radicals. Given that tolerance or resistance to toxic compounds and pathogens can be inherited, female choice should promote the evolution of male ornaments that reliably reveal the status of the bearers' level of oxidative stress. Hence, oxidative stress may be one important agent linking the expression of sexual ornaments to genetic variation in fitness-related traits, thus promoting the evolution of female mate choice and male sexual ornamentation, a controversial issue in evolutionary biology ever since Darwin.     

Observation of a ZZW female in a natural population: implications for avian sex determination

Avian sex determination is chromosomal; however, the underlying mechanisms are not yet understood. There is no conclusive evidence for either of two proposed mechanisms: a dominant genetic switch or a dosage mechanism. No dominant sex-determining gene on the female-specific W chromosome has been found. Birds lack inactivation of one of the Z chromosomes in males, but seem to compensate for a double dose of Z-linked genes by other mechanisms. Recent studies showing female-specific expression of two genes may support an active role of the W chromosome. To resolve the question of avian sex determination the investigation of birds with a 2A: ZZW or 2A: ZO genotype would be decisive. Here, we report the case of an apparent 2A: ZZW great reed warbler (Acrocephalus arundinaceus) female breeding in a natural population, which was detected using Z-linked microsatellites. Our data strongly suggest a role of W-linked genes in avian sex determination.     

Between‐year variation of MHC allele frequencies in great reed warblers: selection or drift?

The major histocompatibility complex (MHC) genes are extremely polymorphic and this variation is assumed to be maintained by balancing selection. Cyclic interactions between pathogens and their hosts could generate such selection, and specific MHC alleles or heterozygosity at certain MHC loci have been shown to confer resistance against particular pathogens. Here we compare the temporal variation in allele frequencies of 23 MHC class I alleles with that of 23 neutral microsatellite markers in adult great reed warblers (a passerine bird) in nine successive cohorts. Overall, the MHC alleles showed a significantly higher variation in allele frequencies between cohorts than the microsatellite alleles, using a multi-variate genetic analysis (amova). The frequency of two specific MHC alleles, A3e (P = 0.046) and B4b (P = 0.0018), varied more between cohorts than expected from random, whereas none of the microsatellite alleles showed fluctuations exceeding the expectation from stochastic variation. These results imply that the variation in MHC allele frequencies between cohorts is not a result of demographic events, but rather an effect of selection favouring different MHC alleles in different years.     

A new approach to study dispersal: immigration of novel alleles reveals female-biased dispersal in great reed warblers

We use the assignment technique and a new approach, the 'novel allele technique', to detect sex-biased dispersal in great reed warblers Acrocephalus arundinaceus. The data set consisted of immigrants and philopatric birds in a semi-isolated population in Sweden scored at 21 microsatellite loci. Fourteen cohorts were represented of which the four earliest were used to define a reference population. Female immigrants had lower assignment probability than males (i.e. were less likely to have been sampled in the reference population), and carried the majority of 'novel alleles' (i.e. alleles observed in the population for the first time). The difference in number of novel alleles between sexes was caused by a strong over-representation of females among the few individuals that carried several novel alleles, and there was a tendency for a corresponding female bias among individuals with low assignment probabilities. Immigrant males had similar or lower reproductive success than females. These results lead us to conclude that important interregional gene flow in great reed warblers depends on relatively few dispersing females, and that the novel allele technique may be a useful complement to the assignment technique when evaluating dispersal patterns from temporally structured data.     

Lost and found: the enigmatic large-billed reed warbler Acrocephalus orinus rediscovered after 139 years

We present compelling evidence of the continued existence of the large-billed reed warbler Acrocephalus orinus , hitherto known only from the unique type specimen collected in NW India 139 years ago. Morphological and genetic analyses of an unusual Acrocephalus warbler mist-netted south-west of Bangkok, Thailand, on 27 March 2006, confirmed its identity as A. orinus , and revealed that it was heterozygous at four out of eight microsatellite markers, indicating the continued existence of a viable population whose breeding and wintering areas are still unknown.     

Secreted PCSK9 downregulates low density lipoprotein receptor through receptor-mediated endocytosis

Proprotein convertase subtilisin/kexin type 9 (PCSK9) is a protease that regulates low density lipoprotein receptor (LDLR) protein levels. The mechanisms of this action, however, remain to be defined. We show here that recombinant human PCSK9 expressed in HEK293 cells was readily secreted into the medium, with the prosegment associated with the C-terminal domain. Secreted PCSK9 mediated cell surface LDLR degradation in a concentration- and time-dependent manner when added to HEK293 cells. Accordingly, cellular LDL uptake was significantly reduced as well. When infused directly into C57B6 mice, purified human PCSK9 substantially reduced hepatic LDLR protein levels and resulted in increased plasma LDL cholesterol. When added to culture medium, fluorescently labeled PCSK9 was endocytosed and displayed endosomal-lysosomal intracellular localization in HepG2 cells, as was demonstrated by colocalization with DiI-LDL. PCSK9 endocytosis was mediated by LDLR as LDLR deficiency (hepatocytes from LDLR null mice), or RNA interference-mediated knockdown of LDLR markedly reduced PCSK9 endocytosis. In addition, RNA interference knockdown of the autosomal recessive hypercholesterolemia (ARH) gene product also significantly reduced PCSK9 endocytosis. Biochemical analysis revealed that the LDLR extracellular domain interacted directly with secreted PCSK9; thus, overexpression of the LDLR extracellular domain was able to attenuate the reduction of cell surface LDLR levels by secreted PCSK9. Together, these results reveal that secreted PCSK9 retains biological activity, is able to bind directly to the LDLR extracellular domain, and undergoes LDLR-ARH-mediated endocytosis, leading to accelerated intracellular degradation of the LDLR.     

Temporal patterns of occurrence and transmission of the blood parasite Haemoproteus payevskyi in the great reed warbler Acrocephalus arundinaceus

We studied the prevalence and intensity of the haemosporidian blood parasite Haemoproteus payevskyi in great reed warblers at Lake Kvismaren (6 years) and Lake Segersj? (3 years) in Sweden. Based on microscopic inspection of slides from 282 adult birds, 20.6% showed infection of H. payevskyi in circulating red blood cells in at least 1 year. For parasite prevalence, there was no difference between years, sex, and age classes. However, parasite intensity was higher in females than in males, and this was most pronounced in 1-year-old birds. Individuals scored to carry parasites in year _n were more likely to show parasite infection year _n + 1 than birds scored to be parasite-free in year _n . None of 99 juvenile birds examined at the breeding site in late summer, 4–9 weeks after hatching, showed infection of H. payevskyi. Parasite intensity in infected adult birds decreased in the course of the breeding season and no new or relapse infections were observed during this period. Thus, our data imply that in the great reed warbler, a long-distance migrant to tropical Africa, transmission of H. payevskyi occurs on wintering sites or at stopover sites during migration.