Sex and environmental sensitivity in blue tit nestlings

In birds and mammals with sexual size dimorphism (SSD), the larger sex is typically more sensitive to adverse environmental conditions, such as food shortage, during ontogeny. However, some recent studies of altricial birds have found that the larger sex is less sensitive, apparently because large size renders an advantage in sibling competition. Still, this effect is not an inevitable outcome of sibling competition, because several studies of other species of altricial birds have found the traditional pattern. We investigated if the sexes differ in environmental sensitivity during ontogeny in the blue tit, a small altricial bird with c. 6% SSD in body mass (males larger than females). We performed a cross-fostering and brood size manipulation experiment during 2 years to investigate if the sexes were differently affected as regards body size (body mass, tarsus and wing length on day 14 after hatching) and pre-fledging survival. We also investigated if the relationship between body size and post-fledging survival differed between the sexes. Pre-fledging mortality was higher in enlarged than in reduced broods, representing poor and good environments, respectively, but the brood size manipulation did not affect the mortality rate of males and females differently. In both years, both males and females were smaller on day 14 after hatching in enlarged as compared to reduced broods. In one of the years, we also found significant Sex × Experiment interactions for body size, such that females were more affected by poor environmental conditions than that of males. Body size was positively correlated with post-fledging survival, but we found no interactive effects of sex and morphological traits on survival. We conclude that in the blue tit, females (the smaller sex) are more sensitive to adverse environmental conditions which, in our study, was manifest in terms of fledgling size. A review of published studies of sex differences in environmental sensitivity in sexually size-dimorphic altricial birds suggests that the smaller sex is more sensitive than the larger sex in species with large brood size and vice versa.     

Tail colour signals performance in blue tit nestlings

Indirect sexual selection arises when reproductive individuals choose their mates based on heritable ornaments that are genetically correlated to fitness. Evidence for genetic associations between ornamental colouration and fitness remains scarce. In this study, we investigate the quantitative genetic relationship between different aspects of tail structural colouration (brightness, hue and UV chroma) and performance (cell‐mediated immunity, body mass and wing length) in blue tit (Cyanistes caeruleus) nestlings. In line with previous studies, we find low heritability for structural colouration and moderate heritability for performance measures. Multivariate animal models show positive genetic correlations between the three measures of performance, indicating quantitative genetic variation for overall performance, and tail brightness and UV chroma, two genetically independent colour measures, are genetically correlated with performance (positively and negatively, respectively). Our results suggest that mate choice based on independent aspects of tail colouration can have fitness payoffs in blue tits and provide support for the indirect benefits hypothesis. However, low heritability of tail structural colouration implies that indirect sexual selection on mate choice for this ornament will be a weak evolutionary force.     

Genetic and environmental variation in immune response of collared flycatcher nestlings

This paper aims at partitioning genetic and environmental contribution to the phenotypic variance in nestling immune function measured with the hypersensitivity test after inoculation with phytohaemagglutinin. A cross-fostering experiment with artificial enlargement of some broods was conducted. Variation in nestling immune response was related to their common origin, which suggests heritable component of cell-mediated immunity. A common rearing environment also explained a significant part of variation. However, deterioration of rearing conditions as simulated by enlargement of brood size did not affect nestling immunocompetence, although it affected nestling body mass. Variation in body mass explained some of the variation in immune response related to rearing environment, which means that growth is more sensitive to the shifts in rearing conditions than the development of immune function. Heritable variation in immune response suggests that there should be potential for selection to operate and the micro evolutionary changes in immunity of flycatcher nestlings are possible.     

Yolk carotenoids increase fledging success in great tit nestlings

Avian mothers can influence offspring phenotype through the deposition of different compounds into eggs, such as antibodies, hormones and antioxidants. The concentration of carotenoids in yolk is larger than in maternal plasma, suggesting an important role of these compounds for offspring development. Since carotenoids have to be acquired from the diet, they may be available in limiting amounts to the mothers. Here, we investigated the role of egg carotenoids for offspring growth by experimentally increasing the concentration of yolk lutein, the main carotenoid in great tit (Parus major) yolk. We subsequently measured body condition, oxidative stress, immune response, plumage colouration and fledging success. Lutein increased body mass soon after hatching and fledging success, but did not affect tarsus length, oxidative stress, immune response and plumage colouration. The higher content of yolk lutein could have increased body mass by reducing oxidative stress caused by high metabolic rates of rapidly growing embryos or by promoting cell differentiation and proliferation. The positive effect of lutein on fledging success seems to be mediated by its influence on body mass 3 days post-hatch, since these two traits were correlated. The finding that our treatment did not affect traits measured later in the nestling period, except for fledging success, suggests that yolk lutein has short-term effects that are essential to increase survival until fledging. Our study shows the positive effect of yolk lutein on offspring survival in the great tit, and therefore suggests an important role of carotenoid-mediated maternal effects.     

Changing climate and the phenological response of great tit and collared flycatcher populations in floodplain forest ecosystems in Central Europe

This study is based on 47 years of observations (1961–2007) on two common bird species, the Great Tit (Parus major) and the Collared Flycatcher (Ficedula albicollis), and a dominant tree species in their habitat, the English Oak (Quercus robur). The study took place at four research sites in the Czech Republic located in full-grown, multi-aged floodplain forests with no forestry management. An increase in air temperature over the evaluated period clearly influenced the length of phenological phases. The full foliage date of English Oak has advanced by 8.7 days during the past 47 years. Great Tit and Collared Flycatcher populations have reacted to the changing climate in the same way, with first laying date and mean laying date advancing by between 6.0 and 9.0 days. In all cases, the trends are highly significant and consistent over all sites. Despite the ongoing shift in phenological stages toward the beginning of the year, the change does not appear to have led to mistiming in the trophic food chain. Overall, this study shows almost identical rates of change in egg laying dates for both bird species in all the floodplain forests studied, and these trends are coherent with those of English Oak and peak herbivorous caterpillar activity.     

Incubation temperature affects growth and energy metabolism in blue tit nestlings

Because the maintenance of proper developmental temperatures during avian incubation is costly to parents, embryos of many species experience pronounced variation in incubation temperature. However, the effects of such temperature variation on nestling development remain relatively unexplored. To investigate this, we artificially incubated wild blue tit (Cyanistes caeruleus L.) clutches at 35.0°, 36.5°, or 38.0°C for two-thirds of the incubation period. We returned clutches to their original nests before hatching and subsequently recorded nestling growth and resting metabolic rate. The length of the incubation period decreased with temperature, whereas hatching success increased. Nestlings from the lowest incubation temperature group had shorter tarsus lengths at 2 weeks of age, but body mass and wing length were not affected by temperature. In addition, nestlings from the lowest temperature group had a significantly higher resting metabolic rate compared with mid- and high-temperature nestlings, which may partly explain observed size differences between the groups. These findings suggest that nest microclimate can influence nestling phenotype, but whether observed differences carry over to later life-history stages remains unknown.     

Marking Activities of the Siberian Roe Deer (Capreolus pygargus,Cervidae) in Central Yakutia

Biology Bulletin - The marking behavior of male Siberian Roe deer in central Yakutia, Siberia is considered in the context of a biological signal field. The optical elements of the Roe deer signal...     

Environmental factors shape cloacal bacterial assemblages in great tit Parus major and blue tit P. caeruleus nestlings

Despite their potential ecological and evolutionary importance, factors shaping the composition of bacterial communities in wild vertebrate populations remain poorly understood. The goal of this study was to examine the relative contributions of environmental factors and genetic factors (e.g. species and common origin) to the variation of cloacal bacterial assemblages in wild bird nestlings. We conducted a partial cross-fostering experiment with two passerine species, the great tit Parus major and the blue tit P. caeruleus, sharing similar habitats and breeding biology. Nestlings of the two species were exchanged four days after hatching and cloacal bacteria were sampled nine days later. The structure of cloacal bacterial communities was determined by Ribosomal Intergenic Spacer Analysis. Our results showed that each nestling displayed a unique bacterial community. Furthermore, nestlings raised in the same nest shared significantly similar bacterial communities. The similarity of bacterial community was higher among heterospecific siblings raised within the same nest than between biological siblings raised in separate nests. Effects of common origin between species could not be detected and, if present, were dominated by nest-based short-term environmental effects. Our results show that growth conditions within nests and individually based endogenous factors have significant effects on cloacal bacteria assemblages and could affect post-fledging condition.     

Increasing selectivity of defense behavior during development of pied flycatcher nestlings

The development of defense reaction was studied in the wildlife and experimentally in 7 broods of altricial pied flycatcher (Ficedula hypoleuca) nestlings. Field studies demonstrated that passive-defense response first appeared on the 4th day of the nest life. It developed from the cessation of begging in young relatively satiated nestlings to characteristic freezing response independent of the level of feeding motivation in older nestlings. Older nestlings also acquire the defense reaction in response to novel visual stimuli. The efficiency of the natural stimulus for defense behavior (species-specific alarm call) nongradually changes during the nest life attaining the 100% level only on the 11th posthatching day. During the initial phase of defense behavior development, the reaction can be induced by different rhythmically organized stimuli. Later it becomes considerably more selective and other rhythmic and acoustic signals become much less effective than the alarm call.     

Immune function and survival of great tit nestlings in relation to growth conditions

Life history theory predicts a trade-off between number and quality of offspring. Reduced quality with increasing brood size may arise from a decrease in body condition or in immunocompetence that would be important in fighting off virulent parasites by immunologically naive offspring. We tested the effect of rearing conditions on immune function of nestling great tits (Parus major) by reducing or increasing broods by two hatchlings. In the middle of the nestling period (on day 8), nestlings from enlarged broods developed lower T cell responses [as measured from the cutaneous swelling reaction to injection with phytohaemagglutinin (PHA)] and tended to have lower total leukocyte and lymphocyte concentrations in their peripheral blood than nestlings from reduced broods. Brood size manipulation affected the PHA response of nestlings most strongly in small clutches, suggesting that nestling immune function was dependent on their parents’ condition, as estimated by original clutch size. Intra-brood differences in nestling mortality were unrelated to immune parameters, but nestlings in broods without mortality had a stronger PHA response, higher concentration of lymphocytes and higher body mass on day 15 than nestlings in broods with mortality. These results support the prediction that the immune function of altricial birds is affected by rearing conditions, and that growth and immune parameters are related to inter-brood differences in nestling survival. Received: 1 February 1999 / Accepted: 19. July 1999     

Polluted environment and cold weather induce laying gaps in great tit and pied flycatcher

We studied the occurrence of laying gaps in free-living populations of the pied flycatcher, Ficedula hypoleuca, and the great tit, Parus major, in a pollution gradient of a copper smelter in south-west Finland. Laying gaps were 2.8 times more common in F. hypoleuca than in P. major. The probability of laying gaps was highest in the heavily polluted zone and lowest in the unpolluted zone for both bird species. Cold weather at the time of laying increased the number of laying gaps in both species, but in P. major this effect was most pronounced in the heavily polluted environment. In the most heavily polluted environment the laying gaps were more likely to occur near the beginning of the laying sequence in both species. The laying gap probability increased with increasing laying date in P. major but not in F. hypoleuca. We suggest that the increased number of laying gaps in the polluted environment results from limited Ca availability and the interference of heavy metals with Ca metabolism in laying females.     

Rearing environment effects on immune defence in blue tit Cyanistes caeruleus nestlings

Rearing conditions may influence ontogeny and functioning of the immune system. Activation of different mechanisms involved in host disease resistance and their internal regulation can be affected by intrinsic and extrinsic factors influencing development. I investigated how rearing environment can influence associations between humoral and cellular constituents of immune defence in nestling blue tits (Cyanistes caeruleus). The ability to mount a cell-mediated immune response was estimated as a hypersensitivity reaction to phytohaemagglutinin, and the ontogeny of humoral immunity was determined by assessing circulating levels of total IgG in 15-day-old nestlings. Heterogeneity in rearing conditions was evoked by placing nest-boxes in areas differing in habitat structural characteristics, and examining natural variation in nest ectoparasite infestations, hatching date, brood size and brood sex-ratio. Habitat characteristics, parasitism and hatching date may shape associations between different components of the immune system in developing birds. I discuss the effects of rearing conditions on the interaction between different arms of the immune system and the implications for understanding negative correlations within the immune system at the individual and brood level.     

Immunocompetence and its costs during development: an experimental study in blue tit nestlings

The allocation into T-cell-mediated immunocompetence was experimentally increased in 68 out of 139 nestlings by supplementary feeding methionine to half of the nestlings in 15 blue tit nests. Methionine-supplemented nestlings had an increased T-cell-mediated immunocompetence, but a reduced growth compared with control siblings. Nestlings that had low initial weights and nestlings that were supplemented with methionine had an increased mortality risk. The investment that nestling blue tits make in immunocompetence appears to be tightly controlled by survival costs paid through a trade-off between immunocompetence and growth.     

Segetal vegetation of Central Yakutia

Seven associations of segetal vegetation are distinguished for the vast arable lands of Central Yakutia. Low species richness and the prevalence of annual weeds are the main features of these communities. Only two dominant speciesSphallerocarpus gracilis andSaussurea amara represent Asian types of areals. Other dominantsFallopia convolvulus, Elytrigia repens, Chenopodium album, Brassica campestris, Lappula squarrosa, Scutellaria galericulata, Thlaspi arvense, Avena fatua andCrepis tectorum are typically cosmopolitan. The phytosociological system of the segetal communities is described.     

Sex-specific development of cell-mediated immunity under experimentally altered rearing conditions in blue tit nestlings

In birds, poor rearing conditions usually have negative effects on T-cell-mediated immune response. However, earlier studies demonstrate that fitness-related traits such as body mass may show sex-specific patterns when subject to alteration of rearing conditions. Therefore, to investigate whether deterioration of rearing conditions influences the development of immune function differently in male and female nestlings, we performed brood size manipulation experiments on blue tit (Parus caeruleus) nestlings. To alter rearing conditions, some broods were increased by three nestlings soon after hatching, while other broods were left non-manipulated. Immune response was assessed as a hypersensitivity reaction to phytohaemagglutinin in 11-day-old nestlings. Additionally, we studied the consequences of brood size manipulation for fledgling body mass and tarsus length. The enlargement of brood size had different effects on the cellular immune responses of male and female nestlings, with males being more negatively affected than their female nest-mates. Sex-specific effects of poor rearing conditions were also recorded for tarsus length, such that tarsus growth was more retarded in female than in male nestlings. We discuss the effects of deterioration of rearing conditions on sex-specific development of cell-mediated immunity with respect to sexual dimorphism of size and developmental strategies in male and female nestlings.     

Cold adaptation in insects of Central Yakutia

The mode of cold hardening was for the first time assessed for 20 insect species living in the extremely cold climate of Yakutia. All insects tested were found to adapt through freeze tolerance, producing ice-nucleating agents that cause the hemolymph to freeze at high subzero temperatures. For the first time ice-nucleating agents were demonstrated in Lepidoptera. Pieris rapae exemplified the possibility of switchover in the survival strategy depending on the climatic conditions.     

Male-biased sex ratio among unhatched eggs in great tit Parus major, blue tit P. caeruleus and collared flycatcher Ficedula albicollis

An increasing number of studies have demonstrated that brood sex ratios are frequently unequal, but the proximate mechanisms underlying this deviation are largely unknown. In the current study we analysed deviation from expected 1:1 sex ratio among dead embryos from unhatched eggs collected from partially unhatched clutches of three passerine bird species. We showed that male embryos were significantly overrepresented among unhatched eggs of great tit Parus major , blue tit P. caeruleus and collared flycatcher Ficedula albicollis . Moreover, the bias in sex ratio differed among species and tended to differ among study years. We discuss several hypotheses to explain the observed male bias among unhatched eggs. We conclude that sex specific embryo mortality may contribute to explain the observed variation in sex ratios in several species of wild birds and that sexing unhatched eggs is important in studies of sex ratio allocation.     

The diet of great tit nestlings: Comparing observation records and stable isotope analyses

The diet of wild animals has been studied using many different strategies, approaches and methods in recent decades. In this regard, stable isotopes analysis (SIA) is becoming a widespread tool, but no study has yet, to our knowledge, compared diet estimations from SIA with direct observations of the diet of passerine nestlings. Accordingly, our aim was to test the predictive power of SIA for this purpose and identify potential confounding factors such as habitat effects. To do this, we compared isotopic signatures of δ¹³C and δ¹⁵N in the feathers of great tit (Parus major) nestlings, and the corresponding estimates of their diet based on stable isotope mixing models, with prey proportions delivered by their parents obtained through video-recordings. Between-nest differences in isotopic signatures of δ¹⁵N were larger than within-nest differences. We found that δ¹⁵N signatures of nestling feathers correlated positively with the proportion of spiders and negatively with the proportion of caterpillars in the nestlings’ diet, the most important prey types. On the other hand, between-nest and within-nest differences in δ¹³C ratios were of similar magnitude and δ¹³C ratios correlated mainly with the proportion of trees surrounding nest-boxes that were Quercus spp. Estimates of diet composition based on mixing models correlated with the observed nestling diet, yet effect sizes were quite low. Although mixing models are commonly used to ascertain diets, our data show that they can provide valuable information on the relative intake of prey types from different trophic levels; but when complex dietary patterns are recorded (e.g. due to the confounding effects of habitat and/or temporal variation) it can be difficult to draw firm conclusions about diet composition.