Feather‐degrading bacteria,uropygial gland size and feather quality in House Sparrows Passer domesticus

Feathers are dead integumentary structures that are prone to damage and thus show gradual degradation over the course of a year. This loss of quality might have negative fitness consequences. Feather‐degrading bacteria are some of the most prevalent feather‐degrading organisms, yet the relationship between feather‐degrading bacteria load and flight feather quality has rarely been assessed. We studied this relationship in free‐living House Sparrows during breeding and non‐breeding annual lifecycle stages. We also considered the size of the uropygial gland, given the antimicrobial function of its secretions, and the effect of body condition. The number of feather holes was positively associated with feather‐degrading bacteria load and was negatively related to uropygial gland size and body condition during the breeding season in both sexes. In the non‐breeding season we found the same relationships, but only in females. The degree of feather wear was unrelated to any of the variables measured during the breeding season, whereas it was negatively associated with uropygial gland size and positively with feather‐degrading bacteria load in the non‐breeding season, but only in females. Our results suggest that feather‐degrading bacteria may induce the formation of feather holes, but play only a minor role in the abrasion of flight feathers.     

Experimental test of a trade‐off between moult and immune response in house sparrows Passer domesticus

A trade‐off between immune system and moulting is predicted in birds, given that both functions compete for resources. However, it is unclear whether such a trade‐off exists during post‐breeding moult. This study tests such a trade‐off in the house sparrow (Passer domesticus). Males injected with an antigen (lipopolysaccharide) significantly moulted slower than sham‐injected males. Moreover, males whose seventh primaries were plucked to simulate moult showed smaller immune response to phytohaemagglutinin than control males, in which seventh primaries were clipped. A trade‐off between moult speed and body mass was also found. The results show a clear trade‐off between moult and immune response in the house sparrow: immune response negatively affected moult and moult negatively affected immune response. These findings suggest that only individuals in good condition may have an efficient moult and simultaneously respond effectively in terms of immunity to pathogens, which could explain how plumage traits honestly indicate parasite resistance in birds.     

Uropygial gland size,feather holes and moult performance in the House Sparrow Passer domesticus

Feather holes represent damage to the plumage of birds and are correlated with delayed moult. Uropygial gland size is negatively correlated with feather holes. Consequently, it was predicted that birds with smaller uropygial glands would have more feather holes, and that this would affect moult performance. I examined this prediction in the House Sparrow Passer domesticus. Individuals with smaller uropygial glands had more feather holes, and those with more feather holes moulted later and faster. Therefore, uropygial gland size seemed to affect moult performance via its effect on feather holes. Uropygial gland size may have a positive effect on plumage quality, through a negative effect on feather holes, and therefore on moult timing and speed.     

Large‐scale spatial variation in feather corticosterone in invasive house sparrows (Passer domesticus) in Mexico is related to climate

Ecologists frequently use physiological tools to understand how organisms cope with their surroundings but rarely at macroecological scales. This study describes spatial variation in corticosterone (CORT) levels in feathers of invasive house sparrows (Passer domesticus) across their range in Mexico and evaluates CORT–climate relationships with a focus on temperature and precipitation. Samples were collected from 49 sites across Mexico. Feather CORT (CORT_f) was measured using methanol‐based extraction and radioimmunoassay. Relationships between CORT_f and spatial and climate variables were examined using simple linear regressions. Ordination was used on climate data, CORT_f was plotted against the resulting axes, and univariate regression trees were used to identify important predictors of CORT_f. Universal kriging interpolation was used to illustrate spatial variation in CORT_f across Mexico. Correlations with ordination axes showed that high CORT_f was associated with low precipitation during the rainy season and low dry season temperatures. Specifically, CORT_f was negatively related to May precipitation and January and July minimum temperatures, and positively related to April deuterium excess and June minimum temperatures. CORT_f was higher in second‐year birds compared to after‐hatch years and after‐second years. House sparrows had higher CORT_f levels in the hot, dry, north‐central region of Mexico, and CORT_f was negatively related to temperature and precipitation. House sparrows molt primarily from August–September but climate conditions throughout the year were important predictors of CORT_f, suggesting that conditions outside of molt can carry over to influence energetics during feather growth. These data suggest that dry conditions are challenging for house sparrows in Mexico, supporting previous work showing that precipitation is an important predictor of broad‐scale CORT variation. This work highlights the utility of CORT_f for evaluating the influence of physiology on current avian range limits; furthermore, these data may allow us to predict future changes in species distributions.     

Circadian entrainment by feeding cycles in house sparrows,Passer domesticus

Summary We studied the potential zeitgeber qualities of periodic food availability on the circadian rhythms of locomotor and feeding activity of house sparrows. The birds were initially held in a LD-cycle of 12:12 h, with food restricted to the light phase. After transfer to constant dim light, the birds remained entrained by the restricted feeding schedule. Following an exposure to food ad libitum conditions, the rhythms could be re-synchronized by the feeding cycle. Shortening of the zeitgeber period to 23.5 h resulted in the loss of entrainment in most birds, whereas a longer zeitgeber period of 25 h re-entrained the rhythms of most birds. Although these results prove that periodic food availability can act as a zeitgeber for the circadian rhythms of house sparrows, several features of our data indicate that restricted feeding is only a weak zeitgeber. The pattern of feeding activity prior to the daily time of food access shown under some experimental conditions suggests that anticipation is due to a positive phase-angle difference of the birds' normal circadian system rather than being caused by a separate pacemaker.     

A taste for novelty in invading house sparrows, Passer domesticus

One of the central questions in invasion biology involves whysome introductions succeed and others fail. Although severalcorrelates of invasion success have been identified, patternsalone cannot identify the mechanisms underlying the invasionprocess. Here, we test the hypothesis that one predictor ofinvasion success, behavioral flexibility, is different betweeninvading and established populations of the same species ofbird. We predicted that neophobia (fear of novelty), a surrogateof behavioral flexibility, would be weaker in an actively invadingpopulation (28 years resident; Colon, Republic of Panama) ofhouse sparrows (Passer domesticus) compared to a populationthat had been resident for more than 150 years (Princeton, NewJersey, USA). To test this hypothesis, we compared latency toconsume novel foods and phobia of novel objects between populationswhen both were kept under similar environmental conditions incaptivity. As predicted, birds from the 150-year-old populationtook significantly longer to approach and consume novel foodsthan birds from the 28-year-old population. Responses to novelobjects were not different between populations however; bothpopulations fed more readily near some novel objects, whichto our knowledge is the first such occurrence in a wild vertebratespecies. Overall, a predilection for trying new foods and beingattracted to novel objects may in part explain how this specieshas so successfully invaded new areas.     

Genetic relatedness in wintering groups of house sparrows (Passer domesticus)

Social behaviour of group-living animals is often influenced by the relatedness of individuals, thus understanding the genetic structure of groups is important for the interpretation of costs and benefits of social interactions. In this study, we investigated genetic relatedness in feeding aggregations of free-living house sparrows ( Passer domesticus ) during the nonbreeding season. This species is a frequent model system for studies of social behaviour (e.g. aggression, social foraging), but we lack adequate information on the kin structure of sparrow flocks. During two winters, we ringed and observed sparrows at feeding stations, and used resightings to identify stable flock-members and to calculate association indices between birds. We genotyped the birds using seven highly polymorphic microsatellite loci, and estimated pairwise relatedness coefficients and relatedness categories (close kin vs. unrelated) by maximum likelihood method. We found that most birds were unrelated to each other in the flocks (mean ± SE relatedness coefficient: 0.06 ± 0.002), although most individuals had at least a few close relatives in their home flock (14.3 ± 0.6% of flock-mates). Pairwise association between individuals was not significantly related to their genetic relatedness. Furthermore, there was no difference between within-flock vs. between-flock relatedness, and birds had similar proportions of close kin within and outside their home flock. Finally, relatedness among members of different flocks was unrelated to the distance between their flocks. Thus, sparrow flocks were not characterized by association of relatives, nevertheless the presence of some close kin may provide opportunity for kin-biased behaviours to evolve.     

Food supplementation affects extrapair paternity in house sparrows (Passer domesticus)

Extrapair paternity (EPP) is common among birds, but the reasonswhy it varies within and among species are less clear. In particular,few studies have experimentally examined how food availabilityinfluences paternity and sexual behavior. We manipulated foodsupply in a nest-box population of house sparrows, Passer domesticus,a colonial passerine with extensive biparental care. Duringthree successive breeding attempts, we changed food availabilityat nest sites and examined behavior and genetic parentage. DNAfingerprinting revealed that the level of EPP within broodswas five times lower in pairs nesting at sites continuouslysupplied with extra food. With extra food, mates spent longertime together at the nest, but this was mainly due to a changein female behavior; females but not males increased total nestattendance. Moreover, we found that individual males did notchange within-pair copulation frequency across treatments, suggestingthat our experiment did not influence male control over fertilizationsthrough copulation behavior. Instead, our study shows that ecologicalfactors can have a strong influence on the time budgets of malesand females, which consequently affects the occurrence of EPP.     

Seasonality in the uropygial gland size and feather mite abundance in house sparrows Passer domesticus: natural covariation and an experiment

The seasonal change, i.e. the marked differences between seasons of low and high productivity, in the abundance of ectosymbionts and the defence intensity of the host against pathogens is a well defined characteristic of temperate zone organisms. Here we investigate the seasonal variation in the uropygial gland size and the abundance of Proctophyllodes feather mites on the wing feathers of house sparrows Passer domesticus in two breeding populations. The size of the uropygial gland varied significantly in male and female house sparrows over the annual cycle. The gland was small during the non‐breeding and mating season, after that it started to grow sharply, reaching its maximum size during breeding. Females had larger gland volumes than males during breeding, and the increase in gland size during breeding was more pronounced in females than in males. The number of feather mites was the lowest during breeding, followed by an increase during moult, and reaching its maximum between the wintering and mating seasons. The absence of a significant relationship between the uropygial gland size and the abundance of feather mites, after controlling for potential confounding variables, supports the view that gland oils do not regulate the number of mites. To investigate further this hypothesis, through a full factorial experimental design we tested the effect of uropygial gland and photoperiod manipulation on the population size and population dynamics of feather mites. The manipulation of uropygial gland had no effect on mites, supporting our observational results. As a result of the experimentally increased day‐length, the abundance of feather mites on wing feathers decreased significantly and more sharply than in the control group, supporting the previous anecdotal evidence about the photosensitivity of these organisms. Using photoperiodic cues, feather mites may respond to seasonal changes that affect their life‐history and population dynamics.     

Constraints on home range behaviour affect nutritional condition in urban house sparrows (Passer domesticus)

In human‐dominated landscapes (semi)natural habitats are typically embedded in tracts of unsuitable habitat. Under such conditions, habitat characteristics and grain size of the surrounding landscape may affect how much food, and at what cost, is available for sedentary species with low home‐range plasticity. Here we combine behavioural radio‐tracking, feather ptilochronology, and landscape analysis to test how nutritional condition varies with home range size in 13 house sparrow [Passer domesticus (Linnaeus, 1758)] populations along an urban gradient. Urban individuals occupied smaller home ranges than conspecifics from rural areas, most distinctly if key cover was highly scattered. In urban plots, patch connectivity, home range sizes, and activity areas were positively correlated, indicating that individual ranging behaviour was related to the spatial distribution of suitable habitat. Urban House sparrows also showed the smallest feather growth bars, which were positively related to home range size at plot level. In contrast, growth bar widths and home range sizes were negatively related in rural populations, whereas in suburban populations, both variables varied independently. We conclude that individuals from progressively more built‐up areas show a restricted ability to adjust their daily ranging behaviour to the scattered distribution of critical resources. This may complement other putative causes of the widespread population decline of urban house sparrows. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 41–50.     

Innate and adaptive immune proteins in the preen gland secretions of male house sparrows

Recent studies have demonstrated that preen oil acts to reduce or eliminate feather-associated bacteria. The mechanisms underlying this antibacterial activity, however, are incompletely understood. In addition to the activity of alcohols (i.e. 3,7-dimethyloctan-1-ol), recent research has suggested that antimicrobial peptides may underlie the antibacterial activity of preen oil. Here, we document the presence of innate and adaptive immune proteins, lysozyme and immunoglobulin Y (IgY), in the preen oil of house sparrows Passer domesticus. We suggest lysozyme functions as an antimicrobial agent, with potentially important impacts against Gram-positive feather degrading bacteria. Furthermore, both lysozyme and IgY likely act in local immune defence of the preen gland, and may also play a role in regulating the local microbiome, with potentially important consequences for chemical communication and signalling. Our findings suggest that the preen gland and its secretions should be considered an integral part of the body's first line of defence against invading infections.     

Genetic structure in insular and mainland populations of house sparrows (Passer domesticus) and their hemosporidian parasites

Small and isolated populations usually exhibit low levels of genetic variability, and thus, they are expected to have a lower capacity to adapt to changes in environmental conditions, such as exposure to pathogens and parasites. Comparing the genetic variability of selectively neutral versus functional loci allows one to assess the evolutionary history of populations and their future evolutionary potential. The genes of the major histocompatibility complex (MHC) control immune recognition of parasites, and their unusually high diversity is genes which is likely driven by parasite‐mediated balancing selection. Here, we examined diversity and differentiation of neutral microsatellite loci and functional MHC class I genes in house sparrows (Passer domesticus), living in six insular and six mainland populations, and we aimed to determine whether their diversity or differentiation correlates with the diversity and the prevalence of infection of hemosporidian parasites. We found that island bird populations tended to have lower neutral genetic variability, whereas MHC variability gene was similar between island and mainland populations. Similarly, island populations tended to show greater genetic differentiation than mainland populations, especially at microsatellite markers. The maintenance of MHC genetic diversity and its less marked structure in the island populations could be attributed to balancing‐selection. The greater MHC differentiation among populations was negatively correlated with similarity in blood parasites (prevalence and diversity of parasite strains) between populations. Even at low prevalence and small geographical scale, haemosporidian parasites might contribute to structure the variability of immune genes among populations of hosts.     

Environmental selection is a main driver of divergence in house sparrows (Passer domesticus) in Romania and Bulgaria

Both neutral and adaptive evolutionary processes can cause population divergence, but their relative contributions remain unclear. We investigated the roles of these processes in population divergence in house sparrows (Passer domesticus) from Romania and Bulgaria, regions characterized by high landscape heterogeneity compared to Western Europe. We asked whether morphological divergence, complemented with genetic data in this human commensal species, was best explained by environmental variation, geographic distance, or landscape resistance—the effort it takes for an individual to disperse from one location to the other—caused by either natural or anthropogenic barriers. Using generalized dissimilarity modeling, a matrix regression technique that fits biotic beta diversity to both environmental predictors and geographic distance, we found that a small set of climate and vegetation variables explained up to ~30% of the observed divergence, whereas geographic and resistance distances played much lesser roles. Our results are consistent with signals of selection on morphological traits and of isolation by adaptation in genetic markers, suggesting that selection by natural environmental conditions shapes population divergence in house sparrows. Our study thus contributes to a growing body of evidence that adaptive evolution may be a major driver of diversification.     

Natural and sexual selection on a plumage signal of status and on morphology in house sparrows,Passer domesticus

Temporal patterns of natural and sexual selection on male badge size and body traits were studied in a population of house sparrows, Passer domesticus. Badge size was a heritable trait as revealed by a significant father-son regression. Survival during autumn dispersal and winter was not related to badge size or body traits in yearling male house sparrows. Badges that signal dominance status were affected positively by directional selection for mating. Adult male house sparrows suffered an opposing selection pressure on badge size during autumn. Contrary to males, female house sparrows did not experience significant directional or stabilizing selection on any body trait. Directional sexual selection on male badge size due to female choice moves male sparrows away from their survival optimum. Opposing directional natural selection on badge size due to autumn mortality caused by predation maintains a stable badge size.     

Absorption and paracellular visualization of fluorescein, a hydrosoluble probe, in intact house sparrows (Passer domesticus)

We describe a method to visualize the cellular location of compounds during absorption by the small intestine in intact animals. First, we employed pharmacokinetic methodology to measure the fractional absorption of sodium fluorescein, a small (MW = 376) water-soluble molecule that is widely used as hydrophilic marker molecule for paracellular permeability studies. Based on the hypothesis that the paracellular pathway acts as a sieve, we predicted that fluorescein absorption would be considerable, but less than that of passively absorbed L-glucose which is a smaller molecule (MW = 180). When the two compounds were gavaged into house sparrows simultaneously, the birds absorbed significantly less fluorescein (42 +/- 8%) than L-glucose (82 +/- 7%), as predicted, and absorptions of the two were correlated as one would predict if they shared the same pathway. We removed intestinal tissue 10 min after gavage with sodium fluorescein and determined the cellular location of the compound's fluorescence using confocal laser microscopy. The fluorescent signal was found primarily in the paracellular space. In contrast, in the same type of experiment using instead the similar-sized fluorescent lipophilic compound rhodamine 123 (MW = 381), most fluorescence appeared inside enterocytes, as expected for a compound that diffuses across the apical membrane. Thus, results from all the experiments are consistent with the hypothesis that hydrophilic fluorescein is absorbed primarily via a paracellular pathway. These methods could be applied to visualize absorption pathways of other compounds in other intact animals.     

Intracellular glucocorticoid receptors in spleen,but not skin,vary seasonally in wild house sparrows (Passer domesticus)

Over the short-term and at physiological doses, acute increases in corticosterone (CORT) titres can enhance immune function. There are predictable seasonal patterns in both circulating CORT and immune function across many animal species, but whether CORT receptor density in immune tissues varies seasonally is currently unknown. Using radioligand binding assays, we examined changes in concentrations of glucocorticoid receptors (GR) and mineralocorticoid receptors (MR) in spleen and skin in wild-caught house sparrows in Massachusetts during six different life-history stages: moult, early winter, late winter, pre-egg-laying, breeding and late breeding. Splenic GR and MR binding were highest during the pre-laying period. This may help animals respond to immune threats through increased lymphocyte proliferation and/or an increase in delayed-type hypersensitivity reactions, both of which CORT can stimulate and in which spleen is involved. A decrease in splenic GR and MR during the late breeding period coincides with low baseline and stress-induced CORT, suggesting immune function in spleen may be relatively CORT-independent during this period. We saw no seasonal patterns in GR or MR in skin, suggesting skin''s response to CORT is modulated primarily via changes in circulating CORT titres and/or via local production of CORT in response to wounding and other noxious stimuli.     

No evidence for differential maternal allocation to offspring in the house sparrow (Passer domesticus)

We tested the differential maternal allocation hypothesis ina population of house sparrows. We experimentally altered theattractiveness of males by treating them with implants filledwith crystalline testosterone (T) or left empty (C). We subsequentlymonitored maternal investment as a function of male hormonaltreatment and the size of the black patch of feathers on thethroat (i.e., the badge), a sexually selected trait. The differentialallocation hypothesis predicts that females should adjust theirinvestment with respect to the benefits they receive by matingwith an attractive male. Given that both circulating levelsof T and badge size are condition-dependent traits, we expectedthat females mated with T males and/or with large-badged malesshould invest more into current reproduction. Contrary to thisprediction, we found no evidence that suggested differentialmaternal allocation in this population of house sparrows. Femaleinvestment in yolk T, yolk mass, clutch size, chick brooding,and feeding was not affected by male hormonal treatment or bymale badge size. As expected, T males invested less into chickbrooding and feeding. More surprisingly, females did not compensatethe reduced paternal contribution to chick feeding. As a consequence,the breeding success of T pairs was largely reduced comparedwith that of C pairs. The absence of differential allocationin a system in which it could have an adaptive role raises thequestion about the possible constraints or overriding factorsoperating on patterns of reproductive investment in this species.