Carotenoid-based colour expression is determined early in nestling life

Carotenoid-based colours are widespread in animals and are used as signals in intra- and interspecific communication. In nestling birds, the carotenoids used for feather pigmentation may derive via three pathways: (1) via maternal transfer to egg yolk; (2) via paternal feeds early after hatching when females are mainly brooding; or (3) via feeds from both parents later in nestling life. We analysed the relative importance of the proposed carotenoid sources in a field experiment on great tit nestlings (Parus major). In a within-brood design we supplemented nestlings with carotenoids shortly after hatching, later on in the nestling life, or with a placebo. We show that the carotenoid-based colour expression of nestlings is modified maximally during the first 6 days after hatching. It reveals that the observed variation in carotenoid-based coloration is based only on mechanisms acting during a short period of time in early nestling life. The experiment further suggests that paternally derived carotenoids are the most important determinants of nestling plumage colou

Carotenoid-based nestling colouration and parental favouritism in the great tit

While elaborate carotenoid-based traits in adult birds may have evolved as honest signals of individual quality in the context of sexual selection or other social interactions, the function of carotenoid-based colours in juveniles is less well understood. We investigated the hypothesis that carotenoid-based nestling colouration has evolved in response to parental preference of intensely coloured offspring during food provisioning. In a field experiment, we manipulated nestling plumage colouration by a carotenoid-supplementation and analysed the parental food provisioning behaviour before feather appearance and at the end of the nestling stage. Carotenoids per se did not influence the nestling's begging behaviour or parental feeding decisions and we found no evidence that carotenoid-based colouration in nestling great tits has a signalling function in parent-offspring interactions. Parents did not discriminate between intensely coloured and control offspring in their food provisioning and in accordance with this finding intensely coloured nestlings were not heavier or larger at the end of the nestling stage. Alternative explanations for the evolution of carotenoid-based colours in nestling birds are discusse

Elevational Changes in Bacterial Microbiota Structure and Diversity in an Arthropod-Disease Vector

Environmental conditions change rapidly along elevational gradients and have been found to affect community composition in macroscopic taxa, with lower diversity typically observed at higher elevations. In contrast, microbial community responses to elevation are still poorly understood. Specifically, the effects of elevation on vector-associated microbiota have not been studied to date, even though the within-vector microbial community is known to influence vector competence for a range of zoonotic pathogens. Here we characterize the structure and diversity of the bacterial microbiota in an important zoonotic disease vector, the sheep tick Ixodes ricinus, along replicated elevational gradient (630–1673 m) in the Swiss Alps. 16S rRNA sequencing of the whole within-tick bacterial microbiota of questing nymphs and adults revealed a decrease in Faith’s phylogenetic microbial alpha diversity with increasing elevation, while beta diversity analyses revealed a lower variation in microbial community composition at higher elevations. We also found a higher microbial diversity later in the season and significant differences in microbial diversity among tick life stages and sexes, with lowest microbial alpha diversity observed in adult females. No associations between tick genetic diversity and bacterial diversity were observed. Our study demonstrates systematic changes in tick bacterial microbiota diversity along elevational gradients. The observed patterns mirror diversity changes along elevational gradients typically observed in macroscopic taxa, and they highlight the key role of environmental factors in shaping within-host microbial communities in ectotherms.Peer reviewe

Bacterial microbiota composition of Ixodes ricinus ticks : the role of environmental variation, tick characteristics and microbial interactions

Peer reviewe

Long-term effects of early parasite exposure on song duration and singing strategy in great tits

Song is a sexually selected trait in many bird species and has been suggested to function as a signal of a male's health and parasite resistance. Here we present an experimental field study on the long-term effects of parasite exposure early in life on adult bird song. We exposed nestling great tits (Parus major) to ectoparasitic hen fleas (Ceratophyllus gallinae) and on their recruitment into the local breeding population assessed the response to a playback of a challenging male in their breeding territory. We show, to our knowledge for the first time in a wild bird population, that parasite exposure early in life affects bird song: song duration of males that were exposed to parasites early in life was reduced by 32% compared with males that grew up in a flea-free environment. Early parasite exposure also significantly reduced the degree of song overlap with the playback, which has been shown to correlate with social status. There was no effect of early parasite exposure on the number of different song types sung or on the latency until the males started the vocal response to the playback. These results suggest that mates or rivals can use song duration and song overlap as proxies for an individual's exposure to parasites early in life. It thereby highlights both the importance of parasites in maintaining honesty of sexually selected traits and the costs of parasitism in terms of reduced attractiveness and competitivenes

Interactions between prenatal maternal effects and posthatching conditions in a wild bird population

Resources and cues provided by the mother before birth are important mediators of developmental plasticity. It has been suggested that the adaptive value of such prenatal maternal effects may depend on the environment encountered by the offspring after birth, and that offspring may perform better when environmental conditions encountered by the mother and the offspring match, than when a mismatch occurs. Here, we test how prenatal maternal effects and postnatal conditions interact in influencing offspring growth and development in wild-living great tits (Parus major) by manipulating food availability experienced by the mother before egg laying, partially cross-fostering nestlings between nests, and manipulating food availability after hatching. We observed significant interaction effects between pre- and postnatal food conditions. Nonsupplemented nestlings reached a similar fledging mass, a trait closely linked to postfledging survival, as food-supplemented nestlings when their biological mother had received extra food during egg laying. It shows that prenatal maternal investment can compensate for growth-limiting conditions after hatching. This effect was sex specific, with daughters benefiting more than sons. Furthermore, food-supplemented nestlings grew largest when their biological mother had not received extra food during egg laying, suggesting that offspring were primed prenatally, possibly through differential egg composition, to use resources more efficiently. However, we found no evidence that offspring performed generally better when pre- and postnatal food conditions matched than when a mismatch occurred. Our results demonstrate the importance of considering the postnatal environment when testing for the ecological and evolutionary consequences of prenatal maternal effects in natural population

A trade-off between reproductive investment and maternal cerebellum size in a precocial bird

Natural selection favours increased investment in reproduction, yet considerable variation in parental investment is observed in natural populations. Life-history theory predicts that this variation is maintained by a trade-off between the benefits of increased reproductive investment and its associated costs for the parents. The nature of these costs of reproduction, however, remains poorly understood. The brain is an energetically highly expensive organ and increased reproductive investment may, therefore, negatively affect brain maintenance. Using artificial selection lines for high and low prenatal maternal investment in a precocial bird, the Japanese quail (Coturnix japonica), we provide experimental evidence for this hypothesis by showing that increased prenatal provisioning negatively affects the size of a particular brain region of the mother, the cerebellum. Our finding suggests that cognitive demands may constrain the evolution of parental investment, and vice versa, contributing to the maintenance of variation in reproductive behaviour in animal populations

The multivariate egg: quantifying within- and among-clutch correlations between maternally derived yolk immunoglobulins and yolk androgens using multivariate mixed models

Egg components are important mediators of prenatal maternal effects in birds and other oviparous species. Because different egg components can have opposite effects on offspring phenotype, selection is expected to favour their mutual adjustment, resulting in a significant covariation between egg components within and/or among clutches. Here we tested for such correlations between maternally derived yolk immunoglobulins and yolk androgens in great tit (Parus major) eggs using a multivariate mixed-model approach. We found no association between yolk immunoglobulins and yolk androgens within clutches, indicating that within clutches the two egg components are deposited independently. Across clutches, however, there was a significant negative relationship between yolk immunoglobulins and yolk androgens, suggesting that selection has co-adjusted their deposition. Furthermore, an experimental manipulation of ectoparasite load affected patterns of covariance among egg components. Yolk immunoglobulins are known to play an important role in nestling immune defence shortly after hatching, whereas yolk androgens, although having growth-enhancing effects under many environmental conditions, can be immunosuppressive. We therefore speculate that variation in the risk of parasitism may play an important role in shaping optimal egg composition and may lead to the observed pattern of yolk immunoglobulin and yolk androgen deposition across clutches. More generally, our case study exemplifies how multivariate mixed-model methodology presents a flexible tool to not only quantify, but also test patterns of (co)variation across different organisational levels and environments, allowing for powerful hypothesis testing in ecophysiology

Artificial selection reveals the role of transcriptional constraints in the maintenance of life history variation

This is the final version. Available on open access from Wiley via the DOI in this recordThe trade‐off between reproduction and self‐maintenance is a cornerstone of life history theory, yet its proximate underpinnings are elusive. Here, we used an artificial selection approach to create replicated lines of Japanese quail (Coturnix japonica) that differ genetically in their reproductive investment. Whole transcriptome sequencing revealed that females from lines selected for high reproductive output show a consistent upregulation of genes associated with reproduction but a simultaneous downregulation of immune genes. Concordant phenotypic differences in immune function (i.e., specific antibody response against keyhole limpet hemocyanin) were observed between the selection lines, even in males who do not provide parental care. Our findings demonstrate the key role of obligate transcriptional constraints in the maintenance of life history variation. These constraints set fundamental limits to productivity and health in natural and domestic animal populations.Universität ZürichKAKENHISchweizerischer Nationalfonds zur Förderung der Wissenschaftlichen Forschun