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Telomere length (TL) early in life often shows high heritability and may predict telomere shortening later in life and life expectancy. Yet, there is limited data about what influences TL and TL change at early developmental stages. It is debated whether early-life TL is determined at conception or shaped by early environmental conditions. Here, we investigate whether TL and telomere shortening are set close to conception. We assortatively paired zebra finches ( Taeniopygia guttata ) based on their TL when nestlings, forming two parental pair groups with either ‘short’ or ‘long’ TL. We then measured TL in the offspring of these pairs at the embryo and nestling stages. In embryos, TL did not differ between offspring from the two parental pair groups. However, in nestlings, particularly in sons, offspring TL matched parental TL. Our results suggest that early-life TL itself is not determined at conception. Instead, telomere shortening rate before and just after hatching appears to determine early postnatal TL. These findings highlight that early development is critical for telomere shortening during early life stages and that it may be a key process underlying the similarity in early-life TL between parents and offspring, potentially affecting telomere dynamics throughout life.

In a broad range of species—including humans—it has been demonstrated that telomere length declines throughout life and that it may be involved in cell and organismal senescence. This potential link to ageing and thus to fitness has triggered recent interest in understanding how variation in telomere length is inherited and maintained. However, previous studies suffer from two main drawbacks that limit the possibility of understanding the relative importance of genetic, parental and environmental influences on telomere length variation. These studies have been based on (i) telomere lengths measured at different time points in different individuals, despite the fact that telomere length changes over life, and (ii) parent–offspring regression techniques, which do not enable differentiation between genetic and parental components of inheritance. To overcome these drawbacks, in our study of a songbird, the great reed warbler, we have analysed telomere length measured early in life in both parents and offspring and applied statistical models (so-called ‘animal models') that are based on long-term pedigree data. Our results showed a significant heritability of telomere length on the maternal but not on the paternal side, and that the mother's age was positively correlated with their offspring's telomere length. Furthermore, the pedigree-based analyses revealed a significant heritability and an equally large maternal effect. Our study demonstrates strong maternal influence on telomere length and future studies now need to elucidate possible underlying factors, including which types of maternal effects are involved.

One group of commonly found parasites in birds, for which fitness consequences and effects on life history traits have been much debated are Haemosporidian blood parasites. In a long term study population of great reed warblers Acrocephalus arundinaceus in Sweden, previous studies have shown that the Haemosporidian blood parasites are in their chronic phase during the breeding season and that the fitness of infected and non‐infected birds are similar. In the present study, we quantified parasite intensity (parasitemia) in 718 adults great reed warblers sampled between 1987 and 1998 for the three most common parasite species; Haemoproteus payevskyi (lineage GRW1), Plasmodium ashfordi (GRW2) and Plasmodium relictum (GRW4). We verified that the q‐PCR method is accurately quantifying Haemoproteus payevskyi (GRW1) as it was highly correlated with the number of parasites seen under microscope. Frequency of mixed infections with two lineages was significantly higher than expected based on the prevalence of each of the three parasite lineages. The mean level of parasitemia was significantly different for the three lineages and individual birds had repeatable parasitemia levels between years. Females tended to have a higher parasitemia than males for all three parasite lineages combined. Females with higher GRW1 parasitemia tended to arrive later in spring to their breeding sites. There was a negative correlation between parasitemia and number of fledged offspring for GRW1, and a tendency for a negative correlation between GRW2 parasitemia and the proportion of recruiting offspring. Overall our results demonstrate that chronic Haemosporidian infections can have slight but significant effects on host life history traits, and therefore may act as important selective agents in wild bird populations.

Maternal effects by which females provide their offspring with non-genetic factors such as hormones, nutrients and antibodies can have an important impact on offspring fitness. In vertebrates, maternal antibodies (matAb) are transferred from the mother, via the placenta, egg yolk or milk during lactation to offspring until they are 2 weeks (birds), 4-10 weeks (rodents) and 9 months (humans) old, respectively. matAb transfer can have direct effects on offspring growth rate in birds and rodents, probably by passively protecting the newborn from common pathogens before their endogenous immune system has matured. Indirect long-term effects of matAb transfer on the offspring's own immunity can be synergistic, if matAb act as antigen templates of the accumulated immunological experience of the mother and educate the newborn's immune system. However, it may also be suppressive if matAb reduce antigen presentation to the newborn resulting in antigen-specific blocking of offspring endogenous immunity. Our aim is to review the mechanisms and direct effects of matAb transfer in vertebrates with an emphasis on birds, outline a framework for research on the long-term effects of matAb on the endogenous immune system of the mature offspring and encourage ecological and evolutionary studies of matAb transfer in non-domesticated animals.

Telomere length and telomere shortening are thought to be critical cellular attributes and processes that are related to an individual's life span and fitness. The general pattern across most taxa is that after birth telomere length gradually decreases with age. Telomere protection and restoration mechanisms are usually assumed to reduce the rate of shortening or at most keep telomere length constant. However, here we have compiled a list of 26 articles showing that there is an increasing number of studies reporting apparent elongation of telomeres (i.e., a net increase in TL from timet to timet+1) often in a considerable proportion of the individuals studied. Moreover, the few studies which have studied telomere elongation in detail show that increases in telomere length are unlikely to be due to measurement error alone. In this article, we argue that episodes of telomere elongation deserve more attention as they could reflect individual strategies to optimise life histories and maximise fitness, which may not be reflected in the overall telomere dynamics patterns. We propose that patterns of telomere (net) elongation may be partly determined by other factors than those causing telomere shortening, and therefore deserve analyses specifically targeted to investigate the occurrence of telomere elongation. We elaborate on two ecological hypotheses that have been proposed to explain patterns of telomere elongation (the ‘excess resources elongation’ and the ‘last resort elongation’ hypothesis) and we discuss the current evidence for (or against) these hypotheses and propose ways to test them. Telomere length can contribute to an individual's life span and fitness. Telomere length gradually decreases with age, however, elongation of telomeres can occur. Telomere elongation could reflect individual strategies to optimise life histories and maximise fitness. We elaborate on the ‘excess resources elongation’ and the ‘last resort elongation’ hypothesis to explain patterns of telomere elongation, reviewing their current support and proposing ways to test them.

Several studies have shown associations between shorter telomere length in blood and weakened immune function, susceptibility to infections, and increased risk of morbidity and mortality. Recently, we have shown that malaria accelerates telomere attrition in blood cells and shortens lifespan in birds. However, the impact of infections on telomere attrition in different body tissues within an individual is unknown. Here, we tested whether malarial infection leads to parallel telomere shortening in blood and tissue samples from different organs. We experimentally infected siskins (Spinus spinus) with the avian malaria parasite Plasmodium ashfordi, and used realtime quantitative polymerase chain reaction (PCR) to measure telomere length in control and experimentally infected siskins. We found that experimentally infected birds showed faster telomere attrition in blood over the course of infection compared with control individuals (repeatedly measured over 105 days post-infection (DPI)). Shorter telomeres were also found in the tissue of all six major organs investigated (liver, lungs, spleen, heart, kidney, and brain) in infected birds compared with controls at 105 DPI. To the best of our knowledge, this is the first study showing that an infectious disease results in synchronous telomere shortening in the blood and tissue cells of internal organs within individuals, implying that the infection induces systemic stress. Our results have far-reaching implications for understanding how the short-term effects of an infection can translate into long-term costs, such as organ dysfunction, degenerative diseases, and ageing.

Dietary carotenoids have been proposed to boost immune system and antioxidant functions in vertebrate animals, but studies aimed at testing these physiological functions of carotenoids have often failed to find support. Here we subject yellow canaries ( Serinus canaria ), which possess high levels of carotenoids in their tissue, and white recessive canaries, which possess a knockdown mutation that results in very low levels of tissue carotenoids, to oxidative and pathogen challenges. Across diverse measures of physiological performance, we detect no differences between carotenoid-rich yellow and carotenoid-deficient white canaries. These results add further challenge to the assumption that carotenoids are directly involved in supporting physiological function in vertebrate animals. While some dietary carotenoids provide indirect benefits as retinoid precursors, our observations suggest that carotenoids themselves may play little to no direct role in key physiological processes in birds. Dietary carotenoids have been proposed to have physiological benefits in addition to contributing to coloration. Here, Koch et al. compare immune and antioxidant functions in yellow, carotenoid-rich vs. white, carotenoid-deficient canaries and find no difference, suggesting a limited physiological role of carotenoids.

Biochemical analyses of blood can decipher physiological conditions of living animals and unravel mechanistic underpinnings of life-history strategies and trade-offs. Yet, researchers in ecology and evolution often face constraints in which methods to apply, not least due to blood volume restrictions or field settings. Here, we test the suitability of a portable biochemical analyser (Zoetis VetScan VS2) for ecological and evolutionary studies that may help solve those problems. Using as little as 80 µl of whole-bird blood from free-living Jackdaws (Corvus monedula) and captive Zebra Finches (Taeniopygia guttata), we show that eight (out of 10) blood analytes show high repeatability after short-term storage (approximately 2 h) and six after 12 h storage time. Handling stress had a clear impact on all except two analytes by 16 min after catching. Finally, six analytes showed consistency within individuals over a period of 30 days, and three even showed individual consistency over a year. Taken together, we conclude that the VetScan VS2 captures biologically relevant variation in blood analytes using just 80 µl of whole blood and, thus, provides valuable physiological measurements of (small) birds sampled in semi-field and field conditions.

The migration patterns of birds breeding at high latitudes have undergone major changes during the Holocene, as species expanded from small refugia following the last glaciation. Unique features of genetic migration programs and species‐specific dispersal patterns have resulted in various levels of migratory connectivity. High migratory connectivity can occur when populations expanding from different refugia maintain historically distinct wintering sites, or when species expanding their breeding ranges maintain a constant migratory vector. Alternatively, species may develop novel routes to nearby wintering sites during range expansion, also leading to high migratory connectivity. Here, we analyse light‐level geolocator and multisensor data logger tracks of great reed warblers Acrocephalus arundinaceus from a population at its eastern range limit in Kazakhstan. We compare their migration routes with published data from five Western Palearctic populations to understand how migration patterns have evolved as the breeding range expanded. Mitochondrial DNA data suggest that Kazakhstan was colonised from the western part of the range, but the logger data show that Kazakh great reed warblers winter in East Africa together with conspecifics from Turkey. This indicates that their migration route did not arise as a simple parallel shift of an unchanged vector‐based programme, but required drastic modifications of the migratory directions to maintain African wintering quarters. A remarkable finding in our study was the detection of a novel spring migration strategy. We found that birds leave the African wintering quarters already in February to spend up to two months at an intermediate staging area in southern Iraq, half‐way to their breeding grounds in Kazakhstan. We call this a two‐step spring migration strategy and discuss the conditions that may promote the evolution of such a behaviour.

Through new tracking techniques, data on timing and routes of migration in long-distance migrant birds are accumulating. However, studies of the consistency of migration of the same individuals between years are still rare in small-sized passerine birds. This type of information is important to understand decisions and migration abilities at the individual level, but also for life history theory, for understanding carry over effects between different annual cycle stages and for conservation. We analysed individual repeatability of migration between years in great reed warblers Acrocephalus arundinaceus; a mediumsized European songbird migrating to sub-Saharan Africa. In seven males, with geolocator data from 2–4 yr per bird, we found low to moderate (non significant) repeatability in timing of migration parameters (R ≤ 0.41), but high (and significant) repeatability for most spatial parameters, i.e. autumn route (R = 0.64) and stopover sites (R = 0.59–0.87) in Europe, and wintering sites (R = 0.77–0.99) in sub-Saharan Africa. This pattern of high spatial but low temporal withinindividual repeatability of migration between years contrasts other tracking studies of migrating birds that generally have found consistency in timing but flexibility in routes. High spatial consistency of migration in the great reed warbler may be due to it being a specialist in wetlands, an unevenly distributed habitat, favouring a strategy of recurrence at previously visited sites. Low temporal repeatability may be caused by large between-year variation in carry-over effects from the breeding season, high flexibility in decision rules during migration or high sensitivity to environmental factors (weather, wind) during migration.