Carotenoid trade-off between parasitic resistance and sexual display: an experimental study in the blackbird (Turdus merula)

Many parasites depress the expression of the carotenoid-based colour displays of their hosts, and it has been hypothesized that animals face a trade-off in carotenoid allocation between immune functions and 'degree of ornamentation'. While numerous correlative studies suggest that parasite infection decreases the intensity of carotenoid-based colour displays, the existence of this trade-off has never been demonstrated experimentally in a host-parasite model. In this study, we used the blackbird (Turdus merula) and Isospora (an intestinal parasite) to assess whether this trade-off does indeed exist. Blackbirds were supplemented with carotenoids while simultaneously being exposed to parasites. Supplemented males circulated more carotenoids in the blood and developed more brightly coloured bills than unsupplemented males. In addition, supplementation slowed down the replication rate of parasites. Supplementation with carotenoids enabled infected birds to maintain their bill coloration, whereas birds that were infected but not supplemented showed reduced bill coloration. At the same time, infection slowed carotenoid assimilation in the blood. Overall, we demonstrated that bill colour reflects a bird's health, and that only males with a carotenoid-rich diet are capable of coping with costs associated with parasitic infection. Carotenoids are thus traded off between host physiological response to parasites and secondary sexual traits. Further investigations are required to determine the physiological mechanisms that govern this trade-off.     

Characterization of 38 microsatellite loci in the European blackbird, Turdus merula (Turdidae, AVES)

We characterized 38 microsatellite loci in the European blackbird, Turdus merula. Thirty-seven loci were identified by testing 242 loci that had been originally isolated in other avian species. One additional locus was isolated from a European blackbird genomic library. All loci were characterized in 20-29 blackbirds from a population in the Czech Republic and displayed between two and 16 alleles, with observed heterozygosity ranging from 0.04 to 1.00. Thirty-seven loci could be assigned a chromosome location in the zebra finch (Taeniopygia guttata) genome based on sequence homology.     

Effects of urbanisation on disease prevalence and age structure in blackbird Turdus merula populations

Despite increasing interest in urban ecology most attention has focussed on describing changes in assemblage composition and structure along urbanisation gradients, whilst relatively little research has focussed on the mechanisms behind these changes. Ecological theory predicts that alterations in biotic interactions are particularly likely to arise, especially with regard to disease risk. Here, we report on differences in prevalence of avian malaria and tick infection and intensity in 11 paired urban and rural blackbird Turdus merula populations from across the western Palearctic. We find large and consistent reductions in tick prevalence and intensity in urban areas. There are also large reductions in the prevalence of avian malaria in many, but not all, urban areas. The proportion of first year birds in urban populations is significantly lower than that in rural ones, and across the more natural rural sites southerly populations contain fewer first years than northern ones. These patterns are expected to arise if survival rates are higher in urban areas, and are negatively correlated with latitude.     

The hemoglobins of the adult blackbird (Turdus merula, Passeriformes). The sequence of the major (HbA) and minor component (HbD)

The blood of the adult blackbird contains one major hemoglobin component (HbA = alpha A2, beta 2, ca. 80%) and one minor one (HbD = alpha D2 beta 2, ca. 20%). The Hb-components were separated by FPLC on a TSK SP-5 PW column, and eluted with a linear NaCl gradient, while the globin chains were purified on a cation exchange (CM-Cellulose). Tryptic peptides from the globin chains were separated by HPLC on an RP-2 Lichrosorb column. The complete amino acid sequence was determined by automatic Edman degradation, using film and gas phase methods. For the alpha A-, alpha D- and beta-chains, peptide alignment was carried out relative to the corresponding chains of the greylag goose (Anser anser). The close phylogenetic relationship between blackbird, tree sparrow and starling is verified by the hemoglobin sequence. The O2-affinities of the major and minor hemoglobin components of the blackbird are not yet known. Thus, the results were interpreted on the basis of primary structure. Substitutions of possible structural significance were examined with the help of molecular graphics/modelling.     

Seasonal changes in egg-mass within and among clutches of birds: general explanations and a field study of the Blackbird Turdus merula

In many populations of birds there is a seasonal change in the mean mass of eggs in the clutch. This might be caused by seasonal changes in the costs of egg-production, or changes in the benefits of laying eggs of different size. In the Blackbird Turdus merula the mean mass of an egg correlated specifically with the air temperature during the period when it was predicted to be undergoing rapid follicular growth. There was no residual effect of date or day-length on egg-mass when statistically controlling for temperature during the period of rapid yolk synthesis, yet temperature still had a significant effect when controlling for date or day- length. Thus the seasonal increase in egg-mass appears to be due to changes in the cost of egg-production, not changes in the benefits of laying larger eggs. However, I could find no effect of food supply during laying on the mean mass of eggs in the clutch, either using an indirect measure of food availability, rainfall, or in a food-supplementation experiment. This could be because females responded to extra food by laying earlier, and probably larger, clutches, rather than by increasing egg-mass. The effect of temperature on egg-mass increased through the laying sequence and there was a small seasonal increase in the mass of the last-laid egg compared with the mean of the other eggs in the clutch. I propose that the mean mass of the last-laid egg relative to the clutch mean, which can characterize a species or population, could evolve in response to the environmentally-caused variance in the mass of the last-laid egg: when the variance is high, the mean may have to be high to avoid producing unviable eggs.     

Long-Term Effects of Chronic Light Pollution on Seasonal Functions of European Blackbirds (Turdus merula)

Light pollution is known to affect important biological functions of wild animals, including daily and annual cycles. However, knowledge about long-term effects of chronic exposure to artificial light at night is still very limited. Here we present data on reproductive physiology, molt and locomotor activity during two-year cycles of European blackbirds (Turdus merula) exposed to either dark nights or 0.3 lux at night. As expected, control birds kept under dark nights exhibited two regular testicular and testosterone cycles during the two-year experiment. Control urban birds developed testes faster than their control rural conspecifics. Conversely, while in the first year blackbirds exposed to light at night showed a normal but earlier gonadal cycle compared to control birds, during the second year the reproductive system did not develop at all: both testicular size and testosterone concentration were at baseline levels in all birds. In addition, molt sequence in light-treated birds was more irregular than in control birds in both years. Analysis of locomotor activity showed that birds were still synchronized to the underlying light-dark cycle. We suggest that the lack of reproductive activity and irregular molt progression were possibly the results of i) birds being stuck in a photorefractory state and/or ii) chronic stress. Our data show that chronic low intensities of light at night can dramatically affect the reproductive system. Future studies are needed in order to investigate if and how urban animals avoid such negative impact and to elucidate the physiological mechanisms behind these profound long-term effects of artificial light at night. Finally we call for collaboration between scientists and policy makers to limit the impact of light pollution on animals and ecosystems.     

Visual pigments, oil droplets, ocular media and cone photoreceptor distribution in two species of passerine bird: the blue tit (Parus caeruleus L.) and the blackbird (Turdus merula L.)

The spectral absorption characteristics of the retinal photoreceptors of the blue tit (Parus caeruleus) and blackbird (Turdus merula) were investigated using microspectrophotometry. The retinae of both species contained rods, double cones and four spectrally distinct types of single cone. Whilst the visual pigments and cone oil droplets in the other receptor types are very similar in both species, the wavelength of maximum sensitivity (λ_max) of long-wavelength-sensitive single and double cone visual pigment occurs at a shorter wavelength (557 nm) in the blackbird than in the blue tit (563 nm). Oil droplets located in the long-wavelength-sensitivesingle cones of both species cut off wavelengths below 570–573 nm, theoretically shifting cone peak spectral sensitivity some 40 nm towards the long-wavelength end of the spectrum. This raises the possibility that the precise λ_max of the long-wavelength-sensitive visual pigment is optimised for the visual function of the double cones. The distribution of cone photoreceptors across the retina, determined using conventional light and fluorescence microscopy, also varies between the two species and may reflect differences in their visual ecology. Accepted: 8 January 2000     

Underlying physiological control of reproduction in urban and forest-dwelling European blackbirds Turdus merula

The development and the continual expansion of urban areas have not only destroyed natural habitats, but also have drastically changed the environmental and ecological conditions of these areas. Consequently, species that have settled in these new man-made ecosystems are exposed to considerable alternations in environmental conditions compared to their 'wild' conspecifics. To understand the impact of human-induced environmental changes on life history events such as reproduction, we compared the timing of the reproductive season and its underlying endocrine control in free-living European blackbirds Turdus merula inhabiting urban and nearby forest areas. Body mass, fat score, gonadal size, luteinizing hormone (LH), testosterone (T), and estradiol (E₂) were measured. Urban blackbirds developed their gonads approximately three weeks earlier than forest birds, whereas the timing of gonadal regression did not differ. There are several factors (e.g. artificial light, temperature, food availability, and social cues) which may have caused the differences in the temporal organization of gonadal growth between the urban and forest-living populations. The advanced gonadal development of urban blackbirds did not coincide with an earlier secretion of reproductive hormones. In contrast, urban males had lower plasma LH and T levels during testicular growth than forest males. Differences in social interactions and environmental conditions may explain the contrast of gonadal development and the timing of hormone secretion between urban and forest blackbirds.     

Vertical transmission of feather lice between adult blackbirds Turdus merula and their nestlings: a lousy perspective

There is limited information about the natural history of the transmission of feather lice (Phthiraptera) from parent birds to their young. This article therefore examines the transmission of 4 species of feather lice from parent blackbirds to their nestlings in an English population, and addresses questions formulated from the perspective of the lice. The lice that disperse onto the several young in the nest were mostly found on the larger chicks, those with higher survival prospects. The lice dispersing to chicks were overwhelmingly nymphs, which cannot be sexed morphologically, and so the prediction that the adult lice dispersing would be disproportionately female, potential founders of a new population, was only supported for the most numerous species, Brueelia merulensis. There was no evidence that louse dispersal to chicks was density dependent and more likely when the parents were more heavily infested. Finally, I predicted that lice might aggregate on female blackbirds, which undertake more brooding, to increase their chance of transmission to nestlings. For 1 louse species, B. merulensis, prevalence, but not louse intensity, was higher on female than male blackbirds. For 2 other louse species, Philopterus turdi and Menacanthus eurysternus, no differences between male and female blackbirds were detected.     

Ultraviolet reflection of berries attracts foraging birds. A laboratory study with redwings (Turdus iliacus) and bilberries (Vaccinium myrtillus)

Unlike humans, birds perceive ultraviolet (UV) light (320 to 400 nm), a waveband which is known to play a role in avian mate choice. However, less attention has been paid to the role of UV light in avian foraging. Some blue, violet and black berries reflect UV light. The colour of berries might be an effective advertisement for avian seed dispersers and indicate the stage of fruit ripeness. We conducted an experiment to test how the UV reflection of berries affects birds'' foraging. Redwings were allowed to choose between UV-reflecting bilberries and rubbed bilberries (UV reduced) in the presence and absence of UV light. We used wild-caught adult and hand-raised juvenile birds to assess possible differences between experienced and naive birds. We found that adult redwings preferred UV-reflecting berries when UV illumination was present, but when UV illumination was absent, they did not distinguish between the two berry types. Our study therefore shows, for the first time, that UV wavelengths are used when birds feed on fruit. However, naive birds showed no preferences, suggesting that age and/or learning may affect frugivore preference for UV reflectance.     

An Experimental Study of Prey-attack Behavior in the European Blackbird Turdus m. merula L

The experiments reported in this paper are designed to investigate the features of insect prey that may influence the orientation of first attack in the European blackbird. Three types of stimulus cues are considered: motion, morphological characters, and color patterns. The ontogeny of prey-handling techniques and the specific manipulatory activities involved in the breaking-up of insect prey are dealt with. The role of learning in the development of preyattack behavior is demonstrated. This study deals only with the preparation of food for selffeeding.     

The reproductive success of Blackbirds Turdus merula in relation to habitat structure and choice of nest site

The reproductive success of a population of Blackbirds Turdus merula occupying farmland and woodland was studied over 3 years to investigate the effects of habitat on breeding success. Territory distribution was patchy in both farmland and woodland; some areas were unoccupied, while other areas were occupied at variable densities. Habitat structure appeared to influence occupation: the index of habitat complexity (cover score) was higher in occupied areas than in unoccupied areas and high-density territories had higher cover scores than low-density territories. However, habitat structure had no significant effect on reproductive success because the cover scores of territories where pairs were successful did not differ significantly from those of territories where there were no successful breeding attempts. There was no evidence of differential mortality rates in adults according to habitat. The height, bulk and exposure of c. 430 nests were measured to determine the effect of nest and nest-site characteristics on reproductive success. Nest exposure was the only feature that differed between successful and failed nests, successful nests being less exposed than failed nests. The major cause of breeding failure was nest predation, but the effect of nest exposure operated only during the laying and incubation period and not during the nestling period. The significance of habitat structure for variation in population densities between habitats is discussed.     

A comparison of infestation patterns by Ixodes ticks in urban and rural populations of the Common Blackbird Turdus merula

Although spatial variation in the patterns of parasite infestations among host populations may have important ecological and epidemiological consequences, the causes underlying such variation are poorly known. In the context of a long-term study on the population biology of Common Blackbirds Turdus merula , we examined the prevalence and intensity of infestation by Ixodes ticks between birds living in rural vs. urban habitats. The overall prevalence of tick infestations was significantly higher in the rural habitat where 74% of individuals ( n  = 130) were infested. This result contrasted markedly with the situation in the urban habitat where less than 2% of individuals ( n  = 360) carried ticks. There was no significant effect of the sex of the host on the intensity or prevalence of tick infestations. There was a significant effect of the age of the host on tick infestations essentially due to the absence of ticks on nestlings. Possible mechanisms responsible for the differences between habitats could include differences in tick survival and/or host resistance towards ticks. Previous studies have shown higher population densities and suggested longer survival for Blackbirds in urban than in rural habitats. Given that ixodid ticks are known to transmit pathogens like Borrelia spp. to wild birds, and that Blackbirds can act as reservoirs for these pathogens, the infection patterns observed in our study area provide a suitable situation to study the interrelations between ticks, Blackbirds and pathogens.